Y^f-t?
HARVARD UNIVERSITY
Library of the
Museum of
Comparative Zoology
m'h.^
BuLLQtln OF THE
Museum of
6 m para five
ogy
Volume 137
1968-1969
'*
HARVARD UNIVERSITY
CAMBRIDGE, MASSACHUSETTS 02138 U.S.A.
CONTENTS
No. 1. The Carabid Beetles of New Guinea. Part III. Harpalinae (Con-
tinued): Perigonini to Pseudomorphini. By Philip J. Darlington.
July, 1968 1
No. 2. Geographic Variation in Anolis distichus Cope ( Lacertilia, Iguan-
idae ) in the Bahama Islands and Hispaniola. By Albert Schwartz.
September, 1968 255
No. 3. Ammonoids of the Late Scythian (Lower Triassic). By Bernhard
Kummel. April, 1969 311
/^K«J
Museum of
Comparative
Zoology
The Carabid Beetles of New Guinea
Part III. Harpalinae (Continued):
Perigonini to Pseudomorphini
p. J. DARLINGTON, JR.
HARVARD UNIVERSITY VOLUME 137, NUMBER 1
CAMBRIDGE, MASSACHUSETTS, U.S.A. JULY 30, 1968
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THE CARABID BEETLES OF NEW GUINEA PART III. HARPALINAE
(CONTINUED): PERIGONINI TO PSEUDOMORPHINI
p. J. DARLINGTON, JR.^
•■ CONTENTS
Introduction to Part III.
Purpose; other parts; acknowledgments _.„ 2
Sources and disposition of material 2
Policies and methods; type examinations;
measurements; drawings 3
Localities 4
Findings 5
Taxonomic section
Tribe Perigonini 5
Perigona 6
Tribe Licinini 14
Badister 15
Physolaesthus 15
Omestcs 16
Dicrochile 16
Microferonia 18
(Tribe Amblystomini) 19
( Aniblystoinits) 19
Tribe Chlaeniini 20
Chlaeniits 1 20
Tribe Oodini 30
Anairichis 31
Oodes 32
Tribe Harpalini 38
Gnathaphanus 4 1
Diuphoromerus 42
Hijphurpax 1 44
Lecanomerus 45
Chydaeiis 47
Platyrnciopus 48
TricJiotichnus 48
Hcirpaloxenus _*._: 59
Lyter ; 63
Coleolissus 64
Hyphacreon 66
Anoplogenius 68
Egadroma 69
Stenolophus 71
Acupalpiis 72
Tribe Anaulacini 76
^ Work supported by National Science Founda-
tion Grant GB-93; see also p. 2.
* Bull. Mus. C
Odoutomasoreus 76
Amuilactis 77
AeplinkUus 77
Cuphoia 78
Tribe Cyclosomini 78
Sarotlirocrepis 78
Tribe Lebiini 80
{ Soniotrichus) 82
Aristolehia 83
Lehia 85
Lach noderma 89
Sinurits QQ
Stenotelus 90
Misceliis 91
Holcoderti.s 94
Minuthodcs 95
Catuscopus 100
Pericalus no
Coptfldera 1 10
Minuphloetis 117
Agonochilu 1I8
Oxyodotitus 122
Moc}ithcni.s 122
( Mochtheroides) 123
DolicJtoctL^ 124
Stricklandia 132
Peliocypas 134
Celacnephes 135
Syntom us 135
Microlestes 135
Apristus 137
( Plochionus) 138
Parena 133
Anch i.sta 139
Eiidynomena 140
Denietrida 140
Phlococarahus 183
Trigunotlwps 184
Nototarus ....'. 185
Anomotarus 186
Tribe Pentagonicini 191
Pentagonica 192
Parascopodes 195
Scopodes 197
Tribe Hexagoniini 202
omp. Zool., 137(1): 1-253, July, 1968 1
Bulletin Museum of Comparative Zoology, Vol. 137, Xo. 1
CONTENTS
Hexagonia 202
Tribe Odacanthini 203
Collium 204
Casnoidca - 207
Bcisi.stich us 208
Clurcncia 209
Dicraspeda 210
Lachnotlionix 214
Etidalia 214
Dobodura 215
Tribe Dryptini 216
Drtjpta - .' 216
Desera 218
Tribe Zuphiini 218
Zuphitim 219
Planctes 220
Tribe Helluodini 222
Pofiona^Iossus- 222
Tribe Ilclluonini 228
Creagris 229
IlcUuoiiidius 229
Ih'lbiopapua 232
HelUiosoma 233
HcUuodcma _. „ 233
Gigadcina 233
Tribe Brathinini 234
Phcro})so])}\its 234
Bracliinus 239
Tribe Pseudomorphini 239
Adelatopus 240
Cn/pt(>cc))halomorpha 242
S))haU(>nH)rpli(i 242
INTRODUCTION TO PART III
Purpose: other parts: acknoidcdiimcnts.
Tliis is the third part of a taxonomic survey
of the beetles of the family Carabidae
(predaeeoiis ground beetles) of the island
of New Guinea.- The present part covers
the tribes of the subfamily Ilarpalinae from
Perigonini through Pseudomorphini. and
thus completes coverage of the famiK
Carabidae in the approximate order of the
Junk-Schenkling Catalogue (Csiki 1932-
f933). I'art IV, which is now being pre-
pared, will be primarily a summary, analy-
- Part I, covering the Cieiiidelinae, Carabinae,
and Flarpalinac llnontili Ptiiostichini (in tlic ordc-r
of the Junk-Schenkhn!4 Catalogue) and Part II,
covering the Agonini, are in the Bulletin nt the
Museum of Comparative Zoology: Part I, in \'ol.
126, No. .3, 1962, pp. 321-564, 4 plates; and Part
II, in Vol. 107, No. 3, 1952, pp. 87-252, 4 plates.
(Because of my special interest in the Agonini,
Part II was written and published lalore Part I.)
sis, and discussion of the New Guinean
carabid fauna as a whole. Among subjects
to be considered are the general nature of
the fauna, its geographic relationships and
origins, its ccologic composition, and its
evolution including specific evolutionary
trends (toward wing atrophy, etc.) and
evolutionary radiations on New Guinea.
However, Part IV will include also a taxo-
nomic supplement to list important new
records of previously recorded species and
to describe a number of additional species
received recently, especially new Agonini
from high altitudes.
I have already acknowledged, in Parts I
and II, aid received from the Guggenheim
Foundation. I have now to acknowledge
also aid received from the National Science
Foundation ( Grant GB-93 ) . which has sup-
ported my work on Carabidae of Ne^^
Guinea in many ways, including publica-
tion of the results.
For meticulous editing and typing of the
manuscript of Part III, I am indebted to
Mrs. Judith Koivumaki, and for the accurate
outline drawings and realistic w atercolors,
to Mrs. Mary Catron.
Sources (iiul disposition of material. Prin-
cipal initial sources of material used in my
\\'ork on New Guinean C^arabidae ha\e been
acknowledged in Part I, page 323, and Part
II, pages 90-91. However, notable addi-
tional material has been received recently.
Most important are thousands of specimens
collected for the Bishop Muscnmi by several
entomologists und(>r the direction of Dr.
J. L. (>ressitt; Mr. Josei Sedlacck and his
wife and son have obtained an especially
large number of Carabidae for the Bishop
Museum. An important collection has hcvu
submitted for stud\ also b\- the Department
ol Agriculture, I'ort Mor(\sb\-. through the
kindness of Mr. J. j. II. S/cnl-I\an\'; this
collection includc\s much matcMial from the
Port Moresby area, and also specimens
from other localities including some Irom
high altitudes. Sent with this collection,
but belonging to him personalK', is a fine-
lot of Carabidae collected bv Dr. U. W.
The Carabid Beetles of New Guinea • Darlington
Hornabrook; this too includes material from
high altitudes. A collection submitted by
the Australian Commonwealth Scientific
and Industrial Research Organization, at
Canlierra, includes specimens from the
Morehead River, on the south coast of
Papua almost opposite the tip of Cape
York; several Australian species not known
elsewhere in New Guinea were found at
this locality. And an interesting collection
has been submitted by the South Australian
Museum, including much material from
Mt. Lamington, Papua.
Because different collections have been
received at different times, and because
different portions of my manuscript have
been finished at different times, I have not
set a single deadline for material included
in the present part of my work. I have
simply used in each case the specimens
available when a given group was studied,
with only a few especially important addi-
tional records interpolated later. Additional
noteworthy records will be included in the
supplement in Part IV, referred to above.
Several of the most productive New
Guinean carabid collectors, whose names
appear many times in the following pages,
are associated with single museums. In
order to save space, I shall cite these col-
lectors without repeating the names of their
museums. The persons in question, and the
museums with which they are associated
and to which their specimens belong, are:
(Ludwig) Biro: Hungarian National
Museum, Budapest
(Miss L. Evelyn) Cheesman: British
Museum
(P. J.) Darlington .(Jr. ): Museum of
Comparative Zoology, Cambridge, Mas-
sachusetts, abbreviated M.C.Z.
(J. L. ) Gressitt: Bishop Museum, Plono-
lulu
Sedlacek(s): Bishop Museum, Honolulu
( Citation of this name in the singular
indicates Mr. Josef Sedlacek; in the
plural, additional or different members
of the Sedlacek family: Marie and/or
J. H. Sedlacek)
(L. J.) Toxopeus: Leiden Museum
Other museums and collections of which
the names are abbreviated are:
American Museum of Natural Ilistorv
(New York): A.M.N.H.
California Academy of Sciences, San
Francisco: Cal. Acad.
Commonwealth Scientific and Industrial
Research Organization, Canberra, Aus-
tralia: C.S.I.R.O.
United States National Museum, Wash-
ington, D. C: U.S.N.M.
Policies and methods; type examinations;
measurements; drawings. My work is
second-stage taxonomy ( see Part I, pp. 328-
330). My methods have been described in
Part I, page 330, and Part II, pages 91ff.
However, I should repeat and stress certain
things. I have tried to be reasonably con-
sistent in preparing descriptions but have
not followed a single model exactly. I
have treated some tribes and some genera
in much more detail than others, the rule
being to give the information that has
seemed worth giving in each case. My
descriptions do follow a basic form but are
flexible in detail. I do not like check-list
taxonomy, in which descriptions are (in
effect) drawn by inserting adjectives in
blank spaces in a standard form. This kind
of taxonomy is easy, but it is likely to be
poor taxonomy. I think it is better to
describe each species individually, following
of course some sort of basic pattern, and if I
state under one species that a character is
striking, I see no reason to state (say)
twenty times under other species that it is
not striking.
Although the present part is consistent
with Parts I and II in general, I have made
a few slight changes of usage to conform
to two publications that have appeared
recently. One is the "Style Manual for
Biological Journals," pulilished in 1960 bv
the American Institute of Biological Sci-
ences, 2000 P St., NW, Washington, D. C,
20016. The other is the revised edition
( 1964 ) of the International Code of Zoologi-
cal Nomenclature. I have in general adopted
BuUetin Museum of Comparative Zoology, Vol 137, No. 1
the details of style suggested by the former,
and ha\'e tried to follow the rules and most
recommendations of the latter. However,
although I have followed the Style Manual
in most ways including most abbreviations,
I have occasionally preferred to follow
Webster's Collegiate Dictionary on points
of general style where I see no reason why
biologists should be different.
References listed under tribes, genera,
and species are limited to items directly
concerned with New Guinea plus selected
items likely to be specially useful to workers
on New Guinean Carabidae.
Type examinations: In the present part of
my work I have indicated what types of
previously described species have been
seen and not seen. I have borrowed for
study a few types in especially difficult
genera in which my work has been in
effect revisionary ( in Perigona, for ex-
ample), but I have not attempted to see or
to borrow types in most cases. There are
two reasons for this. First, I do not think
types should be loaned merely to confirm
identifications in faunal work, especially
when the types come from outside the area
under study, in the present case often from
other islands or from Australia rather than
Irom New Guinea. And second, H. E.
Andrewes saw many of the types in question
and made comparisons with them (see my
Part I, p. 325), and my study of the An-
drewes Collection has enabled me to place
not only his own but also most of the older
Oriental species with reasonable confidence.
I do, however, plan to see many of the
older types, including those in Paris, before
completing Part IV, so that I should be
able to correct errors of identification then.
Measurements. Statements of j^roportions
have been calculated (with a slide rule)
Irom actual measurements made with a
ruled disc in the ocular of a stereoscopic
microscope. Proportions eannol be esti-
mated satisfactorily by eye. When possible,
the proportions are based on measurements
of an average-looking 6 9 . The specimens
thus measured are usually specified in a
paragraph headed Measured specimens, but
this paragraph is omitted under species of
which only one or two indi\iduals are
known. Measurements of length and width
are extremes of all available specimens.
Drawings. My drawings are designed
primarily to show gross form, which is very
difficult to describe in words. Mouthparts,
antennae, and legs are sketched in semi-
diagrammatically. The drawings have been
outlined by Mrs. Mary Catron (usually
with use of a crosslined disc in the ocular
of a stereoscopic microscope), checked by
me ( the checking including measuring and
calculating of proportions ) , and then inked
by Mrs. Catron. I have not tried to figure
all species or even all new ones, but have
tried to show unusual ones and also new
species that are based on only one or two
specimens. I expect to deposit representa-
tive sets of specimens in museums in Lon-
don, Honolulu, Canberra, and elsewhere, as
well as in the continental United States,
and persons working on New Guinean
Carabidae in the future should use m\-
specimens rather than figures of them \\'hich
( like all figures ) are sure to be inadequate.
I have usually not used and therefore not
illustrated genitallic characters. I expect to
discuss this matter — when and how to use
genitallic characters in carabid taxonomy —
in Part IV.
Localities. I plan to include in Part IV
a map showing, as far as possible, all
localities at which C>arabidae ha\e been
obtained in New Guinea. In the meantime
the preliminary map published in Part II,
page 93, shows my own localities, most ol
Miss Cheesman's, and some others, and the
sketch map in Part I, page 326 shows the
route of my collecting on the Bismarck
Range. Also, the Rishoi) Museum has
issued a 19-page "List of New (hiinea
Localities" (to 1966) which gi\es approxi-
mate latitud(\s, longitudes, and altitudes ol
the localities ol Hishop Museum collectors
and ol some other persons. This list is, I
suppose, a\ailal)le on recjuest. I have used
it as a standard h)r spt-Uing ol place names.
The Carabid Beetles of New Guinea • Darlington 5
Certain localities have become especially
important in the course of my work.
Dobodura, Papua, where I collected from
March to July 1944 (see Part I, pp. 325-
326), is by far the best known lowland
locality in New Guinea, for Carabidae.
Wau, in the Morobe District, N-E. N. G., is
by far the best known middle-altitude
locality, thanks principally to the efforts of
the Sedlaceks. And Mt. Wilhelm on the
Bismarck Range, N-E. N. G. (where I
collected), and the Snow Mts., West N. G.
(where Toxopeus collected during the
Netherlands Indian-American (Third Arch-
bold) Expedition of 1938-1939), are the
only venj-higJi-altitude localities well known
for Carabidae. Comprehensive collections
from other localities, especially at high
altitudes, are much needed to show at
what intervals localized species replace
each other on New Guinea. Until this is
known, the total number of species of
Carabidae on the island cannot even be
guessed at closely.
Additional evidence that the label
"Dor(e)y" has been wrongly placed on
many of Wallace's Carabidae that prob-
ably really came from Celebes or the Moluc-
cas is given in the present part of my work:
see, for example, under AmbJystonius (p.
20). For general discussion of this locality
see Part I, pages 330-331. Although many
specimens so labeled evidently did not
come from Dorey, Wallace did go there.
Some of his field notes from there are
quoted under Catascopus in the present
part of my work (p. 102).
Findings. Although analysis and discus-
sion of the New Guinean carabid favma as
a whole will be postponed to Part IV, a
few special points are worth noting now.
Several genera that are chiefly Australian
have been found at high altitudes on New
Guinea, Java, and sometimes other islands
in the Malay Archipelago. These genera
include Mecyclothorax (Part I, pp. 498,
505); Microferonia (present part, p. 18);
and Scopodes (present part, p. 197). One
genus, Chydaeus (p. 47), has been fomid
with an opposite pattern of occurrence, on
the mainland of Asia and at high altitudes
on mountains in the Malay Archipelago
east to New Guinea. However, Bemhidion
and Trechiis have not been found on moun-
tains in New Guinea, although Asiatic stocks
of these genera have reached high moun-
tains farther west in the Malay Archipelago
(Darlington, 1959, Pacific Insects, Vol. 1,
pp. 331-345).
Important evolutionary patterns, of not-
able radiations of Carabidae on New
Guinea, have been found in the Agonini
(Part I) and are described and discussed
for several genera treated in the following
pages. The most striking, in fact exciting,
case is in the lebiine genus Demetrida,
which seems to be in the midst of an
evolutionary explosion. The situation among
these diversely colored carabid beetles in
the mountain rain forests of New Guinea
parallels in some ways the situation among
the birds of paradise in the same forests.
I have seen about 1250 specimens of
Demetrida from New Guinea, representing
apparently 56 species, all new! See discus-
sion under the genus (pp. 142-143) for
further details. Less striking, but neverthe-
less important, radiations of species chiefly
within the confines of New Guinea are
described in Trichotichnus (pp. 48-59),
Catascopus (pp. 101-110), Dolichoctis (pp.
124-132), Anomotarus (pp. 186-191),
Scopodes (pp. 197-202), Dicraspeda (pp.
210-214), and IleUuonidius (pp. 229-232).
TAXONOMIC SECTION
Tribe PERIGONINI
Plattjnini group Perigonae G. H. Horn 1881, Trans.
American Ent. Soc. 9, p. 143.
Perigonini Csiki 1931, Coleop. Cat., Carabidae,
Harpalinae 5, p. 894 ( see for synonymy and
additional references ) .
Jedlicka 1964, Reichenbachia 2, No. 61, pp. 267-
274 ( Oriental forms ) .
Perigonitae Jeannel 1941, Rev. frangaise d'Ent. 8,
p. 137.
Perigonidae Jeannel 1948, Coleop. Carabiques de
la Region Malgache, Part 2, p. 733.
The taxonomic limits of this tribe and its
6 Bulletin Museum of Comparaiive Zoology, Vol. 137, No. 1
relation to other tribes of Carabidae are
doubtful but need not be discussed here.
The only genus of the tribe in New Guinea
is Perigona itself (sensu lato).
Genus PERIGONA Castelnau
Castelnau 1835, Etude Ent., p. 151.
Sloane 1903, Proc. Linn. Soc. New South Wales
28, p. 635.
Andievves 1929, Tijdschrift voor Ent. 72, p. 326
( Suniatran species ) .
Csiki 1931, Coleop. Cat., Carabidae, Haipalinae
5, p. 895 (see for synonymy and additional
references ) .
Jedlicka 1935, Neue Carabiden aus Ostasien, Part
10, pp. 17-19 (Philippine species).
1964, Reichenbachia 2, No. 61, pp. 267-
274 ( Oriental species ) .
Jeannel, see references under tribe, above.
Euriipcri<iona Jeannel 1941, Rev. francaise d'Ent.
8, pp. 138, 149 (new synonymy).
Subgenus Trechicus Leconte 1853, Trans. Ameri-
can Philosophical Soc. 10, p. 386.
Dkigno.sis. Small Tachijs- or Trcchiis-\ike
Carabidae; with usually 2 setae over each
eye; apical segments of palpi rather long,
usually subconical; other technical char-
acters given by Jeannel.
Description. None needed here, except
note that all known New Guinean species
are fully winged.
Tiij)e species. Of Ferigona, P. pallida
Castelnau of Africa; of Eun/perifi,ona, P.
j)rocera Fauvel of Java; of Trechicus, T.
umhripennis Leconte (= Perigona nigriceps,
below ) .
Generic distribution. World-wide in
tropical and warm temperate regions. S(>e
also 4th paragraph of following Notes.
Notes. Ennjperigona Jeannel is based on
Perigona procera Fauvel, a very large
species with maxillary palpi long, slender,
with penultimate segments relatively long.
Perigona rex (below) would go in Euryperi-
goruu if this genus were recognized. How-
ever, "Euri/])crigonu' nitida Jeannel 1941
{= Perigona grandis Jedlicka 1935) of die
Philippines has maxillary palpi relatively
shorter and with pcMiultimate segments
shorter than in procera, and tends to con-
nect the latter with more typical Perigona,
and I do not think generic separation is
advisable.
Jeannel divides Perigona into 2 subgenera
which seem natural and useful. They differ
in arrangement of submarginal elytral
punctures and they differ also in habits:
Perigona sensti stricto occurs (in my ex-
perience) only or mostly on or in logs or
rotting wood; subgenus Trechicus, among
dead leaves or debris on the ground, usuallv
in forest. Perigona (Trechicus) nigriceps
( Dejean ) has extended its ecological range
to include fermenting vegetation and vari-
ous plant materials carried by man, and
has been spread over all the warmer parts
of the world.
Variation of supraocular and lateral
prothoracic setae in this genus is note-
worthy. In rev (below) all these setae are
absent. In P. lata Andrewes of Sumatra
the anterior supraocular and median-lateral
prothoracic setae are absent in both type
and "cotype" in the British Museum. And
in P. astrolahica Csiki the posterior-lateral
prothoracic setae are present or absent, as
described inider this species below.
Species of Perigona are numerous in
tropical Asia and the Malay Archipelago.
Fourteen occur in New Guinea. However,
only 5 (nigriceps and 4 endemic species)
occur in Australia (Darlington, 1964, Psxche
71, pp. 125-129). The New Guinean
Perigona fauna is therefore Oriental in
general naturt> and diversity, and it is Ori-
ental also in relationships of most species,
so far as relationships can be determined.
Farlier ke\s to specic\s of Perigona of the
Malay Archipelago (Andrewes 1929; Jed-
licka 1935, 1964) and Australia (Sloane
1903) have been based principalh' on size
and color, but most ol the 14 New Guinean
species haxc diagnostic structural char-
acters, as the follow ing Key shows. I am
indebted to Dr. Z. Kaszab h)r an opportu-
uit\ to examine the t> pes ot Csikis New
( iuiiicaii species.
""JMic I ol low lug species leeorded Ironi New
Guinea are still unknown to ine, and are
not included in the Key.
The Carabid Beetles of New Guinea
Darlington
SPECIES OF PERIGONA PREVIOUSLY
RECORDED FROM NEW GUINEA BUT
NOT RECOGNIZED FROM DESCRIPTION
Perigona litura (Perroud & Montrousier)
Perroud & Montrousier 1864, Ann. Soc. Linneenne
Lyon 11, p. 72 (Trcchiis).
And'rewes 1929, Tijdschrift voor Ent. 72, p. 372
(in key).
1933, Tijdschrift voor Ent. 76, p. 3C3.
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae
5, p. 897 (see for additional references).
This species was described from New
Caledonia. It is listed by Csiki from sev-
eral islands in the Malay Archipelago, in-
cluding New Guinea, but I cannot find the
source of the New Guinean record. An-
drewes did not know the species. Details
given in the original description, and the
fact that the type(s) occurred in detritus,
suggest that it may be a color form of
nig,nceps.
Perigona subcordata Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 730.
This species was described from the
Kei Islands and is likely to occur in New
Guinea. The size and other details suggest
that it may be an earlier name for astrolabica
Csiki.
Perigona suturaiis Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 728.
The type was collected at Sorong, West
New Guinea, by Beccari and D'Albertis,
and is now in the Genoa Museum. Putzeys'
description does not permit an exact deter-
mination but suggests a small specimen of
astrolabica Csiki or a large one of sub-
cyanescens Putzeys.
Key to Species of Perigona of New Guinea
1. Group of 3 punctures in outer submarsinal
channel of elytron (at % or % of elytral
length) forming a straight line {Perigona
scnsii stricto) 2
- These 3 punctures forming a triangle ( sub-
genus Trechiciis) 9
2. Supraocular and lateral prothoracic setae
absent; very large (9.4-12.4 mm) (p.
8 ) rex
- Two pairs supraocular and usually 2 pairs
lateral prothoracic setae present; size
smaller 3
3. Frontal foveae weak, subobsolete; elytra
each with 2 dorsal punctures, no subapical
puncture above marginal channel; length
c. 2.0-2.5 mm (p. 8) pygmaea
- Frontal foveae short but distinct, margined
externally by weak elevations; elytra with
3 punctures, the 3rd either posteriorly on
disc or subapically above marginal channel;
usually larger 4
4. Elytra with 3rd (posterior) dorsal puncture
on disc, separated from marginal channel
by more than width of latter; if in doubt,
refer here specimens over 4 mm long „ 5
- Elytra with 3rd puncture farther back, just
above edge of marginal channel 7
5. Posterior dorsal elytral punctures less than
Yw of elytral length from apex; length c.
4-6 mm (p. 9) astrolabica
- Posterior dorsal elytral punctures more than
Yxo of elytral length from apex; usually
smaller 6
6. Form normal, moderately broad and de-
pressed; length c. 3.3-4.0 mm (p. 9) _...
sxihcijanescens
- Form narrower, subcylindrical; length 2.6-
3.7 mm (p. 10) papimna
7. Larger, c. 4.5 mm; dark castaneous with
reddish suture and appendages (p. 10 ) - rossi
- Smaller; // approaching rossi in size, form
more depressed and color testaceous ._._ - 8
8. Depressed; nearly uniform testaceous with
head browner but elytra not plagiate; length
slightly over 3 mm (p. 10) livens
- Less depressed; partly testaceous but with
head and much of elytral discs darker;
length under 3 mm (p. 11) plagiata
9. Submarginal channel of elytra behind
puncture-triangle (at % or % of elytral
length ) wide, with bottom flat or convex .. 10
- Submarginal channel behind puncture-tri-
angle very narrow 12
10. Color cither testaceous with dark head and
elytral apices or brownish castaneous with
head slightly darker and suture paler; eyes
large, forming c. right angles with neck;
front and neck with distinct c. isodiametric
reticulation (p. 11) nigriceps
- Color dark castaneous with suture not or
not much paler; eyes variable; microreticu-
lation of head often less distinct, often ( not
always) transverse posteriorly 11
11. Eyes larger; microsculpture of posterior
part of head ( if visible ) not obviously
transverse; length usually c. 3.2—3.6 mm
(rarely smaller) (p. 12) erimae
- Eyes smaller and less prominent; micro-
sculpture of posterior part of head (if
8 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
visible) more trans\'erse; length c. 2.7-
3.3 mm (rarely larger) (p. 12) --_ hidovici
12. Prothorax with sides not strongly sinuate
and posterior angles not denticulate; length
c. 2.8-3.3 nnn (p. 13) lebioides
- Sides of prothorax strongly sinuate or
posterior angles denticulate 13
13. Sides of prothorax strongly sinuate about
% of length before base; (fine) micro-
sculpture present; length c. 3.4 mm (p.
13 ) cordens
- Sides of prothorax nearly straight and con-
verging posteriorly, but posterior angles
aliruptly promini'ntly denticulate; micro-
sculpture absent or nearly so; length c.
3.2-3.4 mm (p. 14) dcntifcr
Perigona is.s.) rex n. sp.
Description. With characters of genus;
form as in Figure 1; very large, broad,
depressed; brownish castaneous, lower sur-
face and legs more reddish; rather shining,
reticulate microsculpture fine, lightly im-
pressed, c. isodiametric on head, slightly
transverse on pronotum and elytra. Head
0.58 and 0.60 width prothorax; mandibles
shorter and more curved than usual in
genus; eyes rather small but prominent,
enclosed behind by genae; antennae with
middle segments c. 1V-' X long as wide;
maxillary palpi slightly shorter than in P.
proccra Fauvel, with apical segments slightly
more conical, and with subapical segments
c. equal length of apical ones; frontal im-
pressions vague; supraocular setae absent;
mentum with a long, triangular tooth.
Prothorax: width length 1.64 and 1.56;
base/apex c. 1.33 and 1.22 (exact measure-
ments imjiossible because basal angles
broadly rounded); lateral setae absent; disc
with fine middle line, other imjiressions
vague. Elytra: width elytra/prothorax 1.20
and 1.24; striae absent or faintly indicated;
each elytron with 2 to 4 dorsal punctures
(variation individual, sometimes xms\in-
metric), anterior puncture larth(>r than
others from suture. Secondary .sexual char-
acters: i front tarsi scarcely dilated but
usually with inconspicuous 2-seriate squa-
mae on first 3 segments below (only near
apex of 1st segment, and soiiietimes missing.
perhaps broken off); S with posterior
femora dentate on upper posterior side near
apex; $ with usually 3, 9 4 or 5 seta-
bearing punctures each side last ventral
segment. Measurements: length 9.4-12.4;
width 3.(S-5.0 mm.
Types. Holotype i (Bishop Mus.) and
1 9 paratype (M.C.Z., Type No. 31,344)
from Sepalakembang, Salawaket Rge., N-E.
N. G., 1920 m, holotype Sept. 11-14 and
paratype Sept. 15, 1956 (E. J. Ford, Jr.);
and the following additional paratvpes.
N-E. N. G.: 2, Wau, Morobe Dist., 1400 m.
Mar. 29, 19^3 (Sedlaceks); 1, same locality,
1650 m, Feb. 23, 1962 (Sedlaceks); 1 L
Feramin, 1200-1500 m, May 2.3-31, 1959
(W. W. Brandt, Bishop Mus.). 1 c? , Okapa
(Busa), [1650-1800 m], Oct. 17, 1964
( Homabrook ) ; 1 c^ , Morae, Kukukuku
[Rge.], E. Highlands, 6000 ft. (c. 1850 m),
Mar. 1, 1964 ( Hornabrook ) . West N. G.:
1 A , Mt. Cyclops, 3500 ft. (1067 m). Mar.
1936 (Cheesman).
Mea.sured specimens. The 6 holotype
and 9 paratype from Sepalakembang.
Notes. This remarkable species would go
in Eiiryperip,ona if the latter were recog-
nized (see discussion under genus). So
far as I know it is unicpie in Feriiiona in loss
of all supraocular and latcMal prothoracic
setae and in the toothed posterior i
femora. It is comparable to P. proccra
Fauvel of Java in size but is broader, and
proccra has the above-mentioned setae and
does not have toothed A femora.
Perigona is.s.) pygmaea Andrewes
Andrewes 1930, Treubia, Supplement 7, pp. 3.34.
345.
Description (for recognition only). A
very small Pcriiiona characterized by wt-ak
frontal sulci and absence of 3rd (subapical)
eKtral punctures; IcMigth c. 2.0-2.5 mm.
Type. Vvo\\\ Bum, collected by Toxo-
peus; now in British Mus. (seen).
Occurrence in New Gtnnea. I*apiia: 4,
Dobodura, Mar.-July lf>44 (Darlington).
N-E. N. G.: 2, lower Busu R., Iluon Pen..
Mav 12 and 17, 1955 (E. O, Wilson. M.C.Z.,
The Carabid Beetles of New Guinea • Darlington 9
1 specimen numbered 1056 ) ; 20, Sattclberg,
1899 ( Biro ) ; 3, Stephansort, Astrolabe Bay,
1898, 1900 (Biro); 5, Aitape, Aug. 1944
(Darlington). West N. G.: 12, Maffin
Bay, Aug. 1944 (Darlington).
Notes. I have this species also from
Leyte and Luzon in the Philippines, and
have examined Andrewes' type from Biiru.
My Dobodura specimens were taken under
bark of rotting logs in rain forest.
Of all New Guinean Peritonei, this
seemed most likely to include short-winged
individuals, but I have examined all speci-
mens listed above, and all are in fact long-
winged.
Perigona is.s.) asfrolabica Csiki
Csiki 1924, Ann. Mus. National Hungary 21, p. 172.
Description. None required here; size,
and number and position of dorsal elytral
punctures are diagnostic, in New Guinea;
length 4.3-6.0 mm except only 3.8 mm in
an apparent dwarf of this species from
Dobodura.
Tijpe(s). From Stephansort, Astrolabe
Bay, N-E. N. G., collected by Biro in 1898;
in Hungarian National Mus. (seen).
Occurrence in New Guinea. Papua: 13,
Dobodura, Mar.-Julv 1944 (Darlington);
1, Kokoda, 1200 ft.' (366 m), June 1933
(Cheesman). N-E. N. G.: holotype + 2,
Stephansort, Astrolabe Bay, 1897 (Biro);
1, Sattelberg, 1899 (Biro); 1, Finschhafen,
Huon Pen., 150 m, Apr. 14, 1964 (Sed-
lacek); 6, Saidor, Gabumi Village, Finis-
terre Rge., June 24-30, July 1-21, 1958
(W. W. Brandt, Bishop Mus.); 1, Wum,
Upper Jimmi Vy., 840 m, July 18, 1955
(Gressitt); 1, Wau, Morobe Dist, 1150 m,
Nov. 7, 1961 (Sedlaceks); 1, same localitv,
1450 m, Feb. 5, 1963 (Sedlacek); 1, same
locality, 1700 m, Feb. 19, 1963 (Sedlacek);
1, Bulolo, "G. Pines," 600 m, Feb. 19, 1962
(Sedlacek). West N. G.: 1, Hollandia,
July-Sept. 1944 (Darlington); 1, same lo-
cality, May 1945 (B. Malkin, U.S.N.M.); 1,
Ifar, Cyclops Mts., 450-500 m, Sept. 9,
1962 (Sedlacek); 1, Maffin Bay, July 8,
1944 (E. S. Ross, California Acad.); 4,
Rattan Camp, 1150 m, Feb.-Mar. 1939
( Toxopeus ) .
Notes. P. astrolahica seems close to but
probably distinct from jacohsoni Andrewes
of Sumatra, in which the suture is red
(usually not red in astrolahica) and the
microreticulation of pronotum and elytra
more transverse. Two more, perhaps re-
lated, apparently undescribed species occur
in Luzon.
All specimens seen from New Guinea
have all usual supraocular and prothoracic
setae (or punctures marking positions of
setae) except that the 4 from Rattan Camp
and the 1 from 1700 m at Wau lack pos-
terior-lateral prothoracic setae. However,
presence or absence of these setae is ap-
parently simple dimorphism, for of 6 speci-
mens from Cape Gloucester, New Britain
(Darlington), 5 lack and 1 has posterior-
lateral setae. Because the distribution of
individuals with and without posterior-
lateral prothoracic setae may be of interest
in the future, I have listed ( above ) all New
Guinean specimens of the species in detail
rather than summarizing the species' oc-
currence.
Perigona is.s.) subcyanescens Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 732.
Csiki 1924, Ann. Mus. National Hungary 21, p. 172.
Andrewes 1930, Treul^a, Supplement 7, p. 334.
Louwerens 1953, Verhandlungen Naturforschenden
Gesellschaft Basel 64, p. 305.
horni Jedlicka 1935, Neue Carabiden aus Ostasien,
Part 10, pp. 18-19 (new synonymy).
Description. None required here. See
preceding Key to Sjjccies for identification;
length ( in New Guinea ) c. 3.3—4.0 mm.
Types. Of subcyanescens, from Andai,
near Dorey, West N. G., collected by
Beccari and D'Albertis, in Genoa Museum.
Of horni, from Imungan, Luzon, in Jed-
licka's collection. (See 2nd paragraph of
following Notes.)
Occurrence in New Guiiiea. Widely dis-
tributed and common on the island: 48
specimens from 13 localities scattered from
Milne Bay to Sansapor, and including
Dobodura and Wau (to 1100 m).
10 BuUetin Museum of Comparative Zoology, Vol. 137, No. 1
Notes. Outside New Guinea, this species
is recorded from West Sumba (Louwerens);
Borneo; Mindanao, Samar, and Luzon in
the Philippines; and doul:)tfully from Burn
(Andrewes); and I have seen specimens
from New Britain and the Solomons.
My identification of mbcyanescem is
based on specimens borrowed from the
Genoa Museum, one marked as compared
with Putzeys' type presumably by Csiki,
and my identification of liorni is based on
comparison with Philippine "cotypes" in
the British Museum.
Perigona is.s.) papuana Csiki
Csiki 1924, Ann. Mus. National Hungary 21, p. 173.
Description. None required here. See
Key to Species, and note subparallel cy-
lindrical form; length 2.6-3.7 mm.
Types. Lectotype (by present designa-
tion) and paratvpe from Stephansort,
Astrolabe Bay, N-E. N. G., 1898 (Biro);
in Hungarian National Mus. The specimen
( sex not determined ) now designated lecto-
type bears the original "Holotypus" label,
although no holotype was specified.
Occurrence in New Guinea. N-E. N. G.:
4 (in addition to the types), Stephansort,
1898 (Biro); 1, lower Busu R., Huon Pen.
May 17, 1955 (E. O. Wilson #1066, M.C.Z.),
in lowland rain forest; 1, Wau, 1300 m,
July 27 (year and collector not given).
Notes. This distinct species seems to be
confined to a limited area on the north side
of N-E. New Guinea.
Perigona is.s.) rossi n. sp.
Description. With characters of genus;
form as in Figure 2, c. as in astrolabica but
slightly more slender and convex; dark
castaneous, suture reddish, appendages red-
dish testaceous; rather shining, reticulate
microsculpture isodiametric on head, in part
transverse on pronotum, more transverse
on elytra. Head 0.70 width prothorax; man-
dibles pointed and slightly curved but not
notably elongate; eyes moderate, narrowly
enclosed behind by genae; antennae monili-
form; palpi with apical segment much
longer than subapical, narrowed and almost
pointed apically; frontal impressions short,
shallow, diverging posteriorly; 2 setae over
each eye. Prothorax: width length 1.43;
base apex 0.90; apex broadly emarginate,
with angles well defined but not advanced
beyond arc of emargination; base emargin-
ate-truncate, with basal angles distinct but
obtuse, slightly blunted; sides broadly
rounded, each with usual 2 setae; disc with
middle line distinct, baso-lateral impressions
weak. Elytra: width elytra prothorax 1.25;
humeri rounded-prominent; apices broadly
but irregularly rounded to obtuse but well
defined sutural angles; striae vaguely in-
dicated; intervals punctulate, 3rd with punc-
tures at c. Vs and % of length and at apex
just above submarginal channel. Secondary
sexual characters: $ unknown; 9 vvith
several setae each side apex last ventral
segment. Measurements: length c. 4.5;
\\'idth 1.7 mm.
Type. Holotype 9 (California Acad.)
from Maffin Bay, West N. G., June 1944
(E. S. Ross); the type is unique.
Notes. This species resembles astrolabica
but differs in details of shape especially of
prothorax, and in position ( nearer apex ) of
posterior elytral punctures.
Perigona is.s.) livens Putzeys
PutzL-ys 1873, Ann. Mus. Ci\ . (ienoa 4, p. 225.
Andrewes 1926, Cat. Philippine Caraliidae, p. 354.
1930, Cat. Indian Insects, Part 18, Ca-
raliidae, p. 265.
Tedlicka 1964, Heielienbacliia 2, No. 61, pp. 268,
270.
Description (lor recognition onh'). A
depressed, pale Teri'^ona s.s. with technical
characters indicated in the preceding Key
to Species; length (in New Guinea) c. 3.3
mm.
Ty])r. DoublfulK from Coromandt>l,
India; via C^haudoir and then Oberthiir
Golls. to Paris Nhis. (not seen).
Occurrence in New Guinea. Papua: 2,
Dobodura, Mar.-July 1944 (Darlington).
Notes. P. liv('i}s is listed b\' Andrewes
The Carabid Beetles of New Guinea
Darlington
11
( 1926 ) from Luzon and Mindanao in the
Philippines, and Andrewes ( 1930 ) indi-
cates that he saw Putzeys' type. I have a
PhiHppine (SE. Bataan) specimen identi-
fied as livens by comparison with An-
drewes' collection. The New Guinean speci-
mens do not match Philippine ones exactly,
but my material is too limited to justify
separating the New Guinean form even as
a subspecies.
Perigona (s.s.) plagiata Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 734.
Andrewes 1930, Cat. Indian Insects, Part 18,
Carabidae, p. 266.
Csiki 1924, Ann. Mus. National Hungary 21, p.
172.
1931, Coleop. Cat., Carabidae, Harpalinae
5, p. 898 ( see for additional synonymy and
references ) .
Jedlicka 1935, Neue Carabiden aus Ostasien, Part
10, p. 18 (in key).
1964, Reichenbachia 2, No. 61, pp. 268,
271.
Van Emden 1937, Stettiner Ent. Zeituns 98, p. .35.
annamita Fauvel 1907, Revue d'Ent. 26, p. 104.
Andrewes 1933, Ann. Mag. Nat. Hist. (10) 11,
p. 110.
Description. A small, brownish testaceous
Perigona s.s. with head and much of elytral
discs darker brown, and with technical
characters as indicated in preceding Key
to Species; length c. 2.2-2.8 mm.
Types. Of plagiata, from Aru and Kei
Islands, collected by Beccari, and from
Andai, West N. G., collected by Beccari
and D'Albertis; in Genoa Mus. Of an-
namita, from Ceylon, Annam, Singapore,
and Andai, West N. G., the specimen(s)
from New Guinea collected by Raffray;
Andrewes (1933) found a "type" in the
Maindron Collection, Paris Mus. Lectotypes
for both plagiata and annamita should be
fixed by the next reviser, after examination
of all the original type material. (Types
not seen. )
Occurrence in Neio Guinea. Common
and widely distributed. I have seen 145
specimens from localities scattered over
most of the length of the island, from
Dobodura to Sansapor; most at low altitudes
but single specimens found at 1100 and
1200 m at Wau.
Notes. Andrewes (1930) records plagi-
ata from a wide range, from SE. Asia and
Japan across the Malay Archipelago to
the Philippines and New Guinea, and
Van Emden lists it from the New Hebrides.
Csiki ( 1924 ) records it from Australia on
the basis of specimens ( which I have seen )
in the Hungarian National Mus., but I think
this is an error (see Darlington 1964,
Psyche 71, p. 125). Perigona rufilabris
(Macleay) of eastern Australia is a similar
but larger species,
Perigona {Trechicus) nigriceps (Dejean)
Dejean 1831, Species General Coleop. 5, p. 44
(Bcmbidium).
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae
5, p. 897 ( see for synonymy and additional
references ) .
Jedlicka 1935, Neue Carabiden aus Ostasien, Part
10, p. 18 (in key).
1964, Reichenbachia 2, No. 61, pp. 268,
270, fig. 2.
Jeannel 1941, Rev. francaise d'Ent. 8, p. 141.
lititra Perroud and Montrousier 1864, Ann. Soc.
Linneenne Lyon 11, p. 72 (Trechus) (new
synonymy ) .
bcccaiii Putzeys 1875, Ann. Mus. Civ. Genoa 7,
p. 732 (new synonymy).
J)iwi Csiki 1924, Ann. Mus. National Hungary
21, p. 173 (new synonymy).
klickai JedHcka 1935, Neue Carabiden aus Ostasien,
Part 10, pp. 18, 19 (new synonymy).
Description (for recognition only). See
preceding Key to Species of Perigona of
Netv Guinea; color either testaceous with
head and apices of elytra darker, or brown-
ish castaneous with suture (and of course
appendages) pale, or intermediate with
elytral disc partly but not entirely clouded;
technical characters include eyes relatively
large and prominent, front isodiametrically
microreticulate, and elytra more conspicu-
ously 3-punctate than usual in the genus,
with posterior puncture usually almost in
line with the others; length c. 2.5-3.0 mm.
Types. Of nigriceps, from North Amer-
ica, sent to Dejean by Leconte; now in
Oberthiir Coll., Paris Mus. Of litiira, from
Kanala, New Caledonia; location of type(s)
12 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
unkno\\'n. Of heccarii, from Sarawak, Bor-
neo, collected by Doria and Beccari; now
in the Genoa Mus. (a lectotype should be
designated by next reviser). Of hiroi, I
now designate as lectot\pe a 9 from Ma-
dang (Friedrich-Wilh.-hafen), N-E. N. G.,
1896 (Biro, Hungarian National Mus.);
this specimen is from Csiki's original series
and is labeled "Holotypus," but the desig-
nation has not been published until now.
Of klichai, from Mt. Makiling, Luzon; in
Andrewes Coll., British Mus. (Types of
hiroi and klickai onlv seen.)
Occurrence in Neic Guinea. Common and
widely distributed at low altitudes: more
than 160 specimens from many localities,
from Milne Bay to "Dorey" and Biak Is.,
and including Dobodura and Wau ( to 1.300
m ).
Notes. P. nigriccps is cosmopolitan,
carried by man to all tropical and warm
temperate regions.
P. lifura, described from New Caledonia
but supposedly widely distributed in the
Malay Archi]ielago, was unknown to An-
drewes. The description fits the dark form
of niii,riceps, and the fact that the type(s)
occurred under vegetable detritus also fits
niiiriceps. (The habitat of niiiriccps is noted
under the genus. ) P. heccarii is another
name lor the dark form of this species ( I
do not consider the dark form worth distin-
guishing by name), and hiroi and klickai,
of which I have seen the types, are also
based on dark examples of ni^riceps.
Perigona {Trechicus) erimae Csiki
Csiki 1924, Ann. Mus. National Hungary 21, p. 173.
Description (for recognition only ). With
characters of Perigona, subgenus Trechicus;
broad, moderately convex; black or castane-
ous with suture usually not paler; eyes
forming c. right angles with neck, but some-
what variable; front with or without (lighth'
impressed) isodianietrie reticulations; pro-
thorax with sides not or slightly sinuate
postci ioily, with angles well defined but
obtuse; elytra not or hiintly striate, with
little or no punctulalion, with subinarginal
channel moderately broad behind puncture-
triangle; length c. .3.2-3.6 mm.
Type. From Erima, Astrolabe Bay, N-E.
N. G., 1896 (Biro); in Hungarian National
Mus. (seen).
Occurrence in Neic Guinea. Thirty-four
specimens from numerous localities in
eastern and central New Guinea, from
Milne Bay and Dobodura to Hollandia and
Cyclops Mts.; not yet found farther west in
New Guinea; most from low altitudes but
reaching 1200 m at Wau.
Notes. Specimens of this species vary
considerablv. The eves varv in size and in
development of genae but usually form
nearly right angles with the neck. Reticulate
microsculpture of the head may be distinct
(but light), or partly obliterated, or c.
absent. And there is some \ariation of
other characters. However, the variation
is not primarily geographic, but occurs at
single localities. I think, but cannot be
quite sure, that only one variable species is
involved.
Csiki's type of erimae is large, with eyes
large and genae slight, and with the front
distinctly reticulate. Proportions of the type
are head 0.83 width prothorax; prothoracic
width length 1.50, base/apex 1.03; width
elytra prothorax 1.53.
Although erimae is known onl\- from New-
Guinea, somewhat similar but apparently
distinct species {andreicesi Jedlicka, arroici
Jedlicka) occur in the Philippines.
Perigona (Trechicus) ludovici Csiki
Csiki 1924, Ann. Mus. National Ilun.uan 21, p. 174.
Description ( for recognition onl\). Form
of PcriiiO)\a, subgenus Trechicus; small;
dark, like dark ni'^riceps but suture not or
not conspicuously reddish; head with e\'es
smaller than in niil.rice))s. Iront less dis-
tinctK' reticulat(> and with rtiieulations
more trans\'erse espeeialK posteriorly;
elytra with .3rd (posti'rior) punctures nearer
sutm(>; length c. 2.7-.3.3 mm.
T\i])cs. Lectotype (present designation)
from Mt. llansemann, Astrolabe Bay, N-E.
N. (;., 1901 (Biro, Tbnigarian National
The Carabid Beetles of New Guinea • Darlington
13
Mus. ), and 8 additional original (co) types,
2 with same data as lectotype and 6 from
Madang (Friedrich-Wilh.-hafen), 1900 and
1901 (Biro) (seen).
Occurrence in Neiv Guinea. Seventy-five
specimens from numerous localities over
almost the whole length of New Guinea
(Milne Bay and Dobodura to the Vogel-
kop); at low altitudes, none above 550 m.
Notes. P. ludovici is compared with
nigriccps in the preceding Description. P.
ludovici is in fact closer to erimac but has
the head narrower, eyes relatively smaller,
and microreticulation of posterior part of
head usually more transverse. Also, ludovici
averages smaller than erimae, although
measurements of length overlap: ludovici,
c. 2.7-3.3; erimae, c. 3.2-3.7 mm. (Csiki
gives 2.5-2.8 mm for ludovici and 3.5 mm
for erimae. ) Nevertheless, these species are
very similar and some individuals are dif-
ficult to place. Proportions of the lectotype
of ludovici are head 0.76 width prothorax;
prothoracic width/length 1.49, base/apex
1.08; width elytra/prothorax 1.59.
Both erimae and ludovici live among dead
leaves on the ground in rain forest.
Perigona (Trechicus) lebioides Csiki
Csiki 1924, Ann. Mus. National Hungary 21, p. 174.
Description ( for recognition only ) . Form
of small, very broad, moderately convex
Perii^ona, subgenus Trechicus; castaneous
with suture not or only faintly reddish;
prothorax with sides not strongly sinuate
and not denticulate posteriorly; elytra with
submarginal depressed space very narrow
behind puncture-triangle; elytra faintly or
irregularly striate, not or not much punc-
tulate; length 2.8-3.3 mm.
Types. I now designate as lectotype the
specimen marked "Holotypus" by Csiki. It
is from Erima, Astrolabe Bay, N-E. N. G.,
1896 (Biro) in Hungarian National Mus.
(seen). Seven paratvpes are from Sattel-
berg, N-E. N. G., 1899 (Biro). (Two addi-
tional specimens labeled as paratypes of
lebioides, from Simbang, Huon Gulf, Biro,
in Plungarian National Museum, are not
lebioides but erimae. )
Occurrence in Neiv Guinea. Sixty-one
specimens (including 44 from Dobodura)
from localities in all 3 political divisions of
New Guinea; most at low altitudes but 1
from Sibil, Star Rge., at 1260 m ( Leiden
Mus.).
Notes. This, like the other small Perigona
of subgenus Trechicus that occur in New
Guinea, lives among dead leaves on the
floor of rain forest. Biro presumably col-
lected the types by sifting. I took mine
by throwing raked-up leaves and leaf mold
into still water, and picking up the beetles
as they came to the surface.
Perigona {Trechicus) cordens n. sp.
Description. With characters of Perigona,
subgenus Trechicus; form broad, rather
convex; black or castaneous, suture not or
not much paler, elytra subiridescent, ap-
pendages reddish testaceous; reticulate
microsculpture faint, not clearly visible at
c. 100 X l)ut apparently isodiametric on
front, somewhat transverse posteriorly on
head, fine and strongly transverse on
pronotum and elytra. Head 0.80 and 0.79
width prothorax; eyes rather large, forming
c. right angles with neck, mandibles average
for genus; antennae with middle segments c.
IVi X long as wide; front with impressions
irregular but distinct, margined externally
by short elevations. Prothorax cordate;
width length 1.47 and 1.47; base apex 0.98
and 0.97; sides strongly sinuate about Vs
from base; posterior angles nearly right but
blunted; disc with fine middle line, shallow
poorly defined baso-lateral impressions.
Elytra short, wide; width elytra prothorax
1.64 and 1.66; submarginal impressed space
very narrow^ behind puncture-triangle; each
elytron with parts of at least 6 striae, inner
ones moderately impressed and irregular
or vaguely punctate; intervals not distinctly
punctulate, 3rd 3-punctate. Secondary sex-
ual characters normal: c5 front tarsi with
3 segments (only apex of 1st) narrowly 2-
seriately squamulose; 6 with 2, 9 c. 4
14 BuUetin Museum of Comparative Zoology, Vol. 137, No. 1
setae at apex last ventral segment. Mea-
surements: length c. 3.4; width 1.5-1.6 mm.
Types. Holotype $ (M.C.Z., Type No.
31,345) and 3 paratypes (broken <^ , 9 $ )
all from Dobodura, Papua, Mar.-July 1944
( Darlington ) .
Measured speeimens. The 6 holotype
and 1 9 paratype.
Notes. This species occurred in leal
litter in rain forest, with erimae, ludoviei,
and lehioides, from all of which cordens is
immediately distinguishable by its strongly
cordate prothorax.
Perigono (Trechicus) denfifer n. sp.
Description. With characters of Perigona,
subgenus Trechicus; form as in Figure 3;
broad, moderately convex; reddish castane-
ous with suture slightly paler, appendages
reddish testaceous; shining, not iridescent,
microsculptme absent or nearly so. Head
0.78 and 0.78 width prothorax; mandibles
slender, pointed, weakly arcuate near apex;
eyes moderately large but less prominent
than usual, forming obtuse angles with
neck; antennae with middle segments c.
Vri X long as wide; front with slight median
puncture and distinct short anterior frontal
impressions. Prothorax broadly subcordate,
very wide anteriorly; width/length 1.40
and 1.45; base/apex 0.95 and 0.88; sides
weakly rounded, strongly converging pos-
teriorly almost to base, then abruptly sinu-
ate with basal angl(\s right-denticulate; disc
with usual middle line and transverse im-
pressions, basal transverse impression sub-
foveate at middle and running into slightly
dccjier but poorly defined baso-lateral im-
pressions. FJijtra wide; width elytra pro-
thorax r. 1.61 and 1.60 (exact measurement
impossible because elytra spread in both
specimens); submarginal impressed space
very narrow behind puncture-triangle; 6
abbrc-viated striae on each elytron, inner on(\s
impressed, all plainU' punctate; int(>r\als
not punclulate, 3rd 3-puuctate. Secondary
.sexual characters as in jireceding species
(cordens). Measuremenl.s: length r. 3.2-
3.4; width r. 1.4-1.5 mm.
Types. Holotype S (M.C.Z., Type No.
31,346) and 1 9 paratype both from Milne
Bay, Papua, Dec. 194.3 (Darlington).
Notes. The form of prothorax, absence of
microsculpture, and impressed punctate
elytral striae are diagnostic of this species.
Tribe LiCININI
Sloane 1898, Proc. Linnean Soc. New South Wales
23, pp. 487 ff . ( Australian genera ) .
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae
.5, p. 899.
Ball 1959, Mem. American Ent. Soc, No. 19, p.
5 ( see for synonymy and additional references ) .
Most Licinini, including all those known
from New Guinea, have the labrum and
usually also the clypeus deeply emarginate,
the labrum often so deeply so as to appear
2-lobed. This alone is almost a sufficient
recognition character of the tribe, in New
Guinea. Other diagnostic characters are
discussed by Ball ( 1959, pp. 5-8).
Licinines are nearly world-wide in distri-
bution but are relatively few in Central
and South America and relatively numerous
in Australia: about 10 genera, including
some that are probably primitive or relict,
occur in Australia. Five genera occur in
New Guinea: Badister, which is \\idel>
distributed in other parts of the world;
Omestes, a monotypic genus confined to the
eastern part of the Mala\^ Archipelago;
and Physolaesthus, Dicrochilc. and Micro-
feronia, which are primarih Australian.
Thrc^e species of Dicrochile and one of each
of the other genera are known in N(^w
Guinea. All the New Guinean species are
winged, except Microferonia haro.
The following Key is based on Balks
(1959, p. 11) key to Oriental licininc^
genera.
Kky to Gfnfha ok Licinini ok Xknv Ciinka
1. One mandible deeply notched alioxe, with a
prominent boss beliind tiir iiotrh 2
Neither mandible notched as described 4
2. I.cil mandible notched; only basal seginent
oi antenna glabrous (p. 15) ._. Badister
- \\\si}\{ mandible notchecb each antciuKi w itii
3 segments glabrous 3
3. Smaller (c. 5 mm); elytra not spined (p.
15) Phtjsolacsthus
The Carabid Beetles of New Guinea • Darlington
15
- Larger (c. 11-15 mm); elytra with short
apical spines (p. 16) Omcstc.s
4. Form Agonu)n-\\\.e; mandibles blunt at apex
(p. 16) — Dicrochile
- Form elongate-oval with very small head;
mandibles ( at least the right one ) con-
spicuously 2-dentate at apex, with upper
tooth large, acute (p. 18) Miciofeiouia
Genus BADISTER Clairville
Anonymous [Clairville] 1806, Entomologie Hel-
vetique 2, p. 90.
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae
5, p. 901 (see for synonymy and additional
references ) .
Jeannel 1942, Faune de France, Coleop. Carabiques,
Part 2, p. 1000.
Louwerens 1956, Treubia 23, p. 236 (key to
Indonesian species ) .
Ball 1959, Mem. American Ent. Soc, No. 16, pp.
189-191.
Diagnosis. See preceding Key.
Description. None required here.
Type species. Carabus bipustidatus Fab-
ricius, of Europe, etc.
Generic distribution. Temperate and
tropical Eurasia, the Malay Archipelago,
and eastern Australia; Africa and Mada-
gascar; North America and some West
Indies, but not South America.
Notes. See Jeannel (1942) and Ball
( 1959 ) for further information on this
widely distributed genus.
Badisfer (Baudia) sundaicus Andrewes
Andrewes 1926, Ann. Mag. Nat. Hist. (9) IS, p.
275.
Louwerens 1956, Treubia 23, p. 236.
Description. See Andrewes (1926); length
c. 4.0-4.5 mm.
Type. From Soekaboemi, Java; in
Andrewes Coll., British Mus. (seen).
Occurrence in New Guinea. Papua: 4,
Dobodura, Mar.-July 1944 (Darlington).
N-E. N. G.: 1, Maprik, Sepik Dist., 150 m,
Dec. 29, 1959-Jan. 17, 1960 (T. C. Maa,
Bishop Mus.). West N. G.: 2, Hollandia,
July-Sept. 1944 (Darlington).
Notes. I tentatively identify as sundaicus
specimens from Siani and the Malay Pen.
(British Mus.); Sumatra; Java; Luzon and
Leyte in the Philippines; Morotai Is. in
the Moluccas; New Guinea (listed above);
New Britain; and widely scattered locali-
ties in eastern Australia. Specimens from
all these places have the mandibular and
antennal characters indicated in the pre-
ceding Key to Genera. However, variation
is obvious, and further study may show
that more than one species is involved.
Specimens of this and related species
that I have collected were usually among
dead leaves and vegetation on the ground
in very wet places by standing (not running)
water.
Genus PHYSOLAESTHUS Chaudoir
Chaudoir 1850, Bull. Soc. Nat. Moscow 23, Part
1, No. 2, p. 411.
Diagnosis. See preceding Key to Genera.
Description. See Chaudoir (1850), and
following Notes.
Type species. P. australis Chaudoir, of
Australia.
Generic distribution. Primarily Austra-
lia; one species described from New
Zealand; and the following species (if
correctly assigned) on New Guinea, Java,
and the Philippines.
Notes. I have not been able to identify
australis in the Australian material before
me. Chaudoir does not describe its antennal
pubescence but states that the right man-
dible is tuberculate, and this character is
always associated with 3 antennal segments
glabrous, among Australian licinines known
to me. Whether the following species is
really a Fhysolaesthus and how this genus
is related to Badistcr will have to be decided
by future revisers.
Fhysolaesthus caviceps (Andrewes)
Badistcr caviceps Andrewes 1936, Ann. Mag. Nat.
Hist. (10) 17, p. 312
Louwerens 1956, Treubia 23, p. 236.
Description. See Andrewes, and my
Figure 4; length c. 5 mm.
Type. A $ from Toeloengagoeng, Java;
in Andrewes Coll., British Mus. (seen).
Occurrence in New Guinea. West N. G.:
4, all from Wissel Lakes area, as follows:
16 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
1, Itouda, Kamo Vy., 1500 m, Aug. 12,
1955 (Gressitt), in light trap; 1, Lake
Paniai, 1570 m, Aug. 28, 1939 (H. Boschma,
Leiden Mus.); 2, Enarotadi. LSOO m, Aug.
1, 1962 (Sedlacek).
Notes. I have seen specimens with the
characters of caviceps from Java and
Luzon as well as New Guinea but am not
sure whether they represent one species or
two or more related species. Except for
the different mandibles and antennae, this
species is remarkably similar to Badisfcr
sundaicus (above), and I think the habitats
of the two species are similar, judging
from what I have seen of them in the
Philippines.
Genus OMESTES Andrewes
Andrewes 1933, Treubia 14, p. 276.
Diafinosis. See preceding Key to Genera.
Description. See Andrewes.
Type species. Omestes torta Andrewes,
below.
Generic distribution. Same as that of O.
torta, below.
Notes. I suspect that Omestes torta may
prove to be only a large, specialized
(spined) Physokiestlius, but I shall leave a
decision about this to future revisers,
Omestes torta Andrewes
Andrewes 1933, Treuljia 14, p. 277.
Louvverens 195(-), Treul)ia 23, p. 224.
Description. See Andrewes, and m>
Figure 5; length 11-14 mm.
Type. A £ from Sangi Is.; in Andrewes
Goll., British Mus. (seen).
Occurrence in New Gttinea. Papua: 1,
Dobodura, Mar.-July 1944 (Darlington); 1,
Milne Bay, Dec. 1943 (Darlington); 2,
Kiunga, Fly R., Aug. 14-17, 1957 (W. W.
Brandt, Bishop Mus.); 1, Darn Is., Mar. 16-
31, 1936 (Archbold Expedition, A.M.N.H.);
1, Woodlark Is. (Murua), Kuhunadau ITill,
Apr. 16-22, 1957 (W. W. lirandl. Bishop
Mus.). West N. C;.: 19, Mollandia, Julv-
Sept. 1944 (Darlington); 1, Maffen, Tor R.
(mouth), 4 km E. of Hollandia, JuK 2.
1959 (T. G. Maa, Bishop Mus.), at light;
1, Bernhard Gamp, 50 m, Apr. 12, 1939
( Toxopeus ) .
Notes. Omestes torta is now known from
New Guinea, the Moluccas ( Ilalmahera
and Morotai), Celebes, the Sangi and
Talaud Islands, and the Philippines
(Leyte). My material is not sufficient to
show details of geographic variation. The
insect lives among dead leaves and vege-
tation on the ground in deep swamps.
Genus DICROCHILE Guerin
Guerin 1<S46, Ann. Soc. Ent. France (2) 4, Bnll.
p. cm.
Sloane 1923, Proc. Linnean Soc. New South Wales
48, pp. 35-36 (key to Australian species).
Csiki 1931, Colcop. Cat., Carabidae, Harpalinae 5,
p. 921 ( see for synonym\- and additional refer-
ences ) .
Diagnosis. See preceding Key to Genera.
Description. None required here.
Ty})c species. Presumabh- Dicrochilc
fabrii Guerin, of New Zealand. (I do not
wish to designate a type species. If no
formal designation has been made, it should
be left to the next reviser. )
Generic distribution. New Zealand, Aus-
tralia, New (Guinea, Moluccas (Obi Is.),
Solomons ( a probably undescribed species
near alternans from Bougainville), New
(Caledonia.
Notes. All species of this genus that I
know, in Australia as well as New Ciuinea,
arc \\'inged. Most of them live in swamps
or other wet places, but alternans (de-
scribed below) is a nicsophile.
Kkv k) Species oi- Dickociiilk ok Xeav Guinea
1. I']l\tra w'itli atiitc tcctli or sliort spini's at
sutnral and outer-apical ant^lcs; dorsal elytral
intcr\als ccjnal or ncarK so (p. 10) acuta
- Elytra not toothed or spined; dorsal ehtral
inter\als uncciual _ ._ 2
2. I'^ront of head normalK' con\e\; smaller,
length 11.5-12.5 nun (p. 17) altcntans
— I'roiit ol licad sliyhtK depressed; hir^cr,
length 13. .5-14. 5 nnn ( [i. IS) tiro
Dicrochile acuta n. sp.
Description. Form (Fig. 6) ol ALi,()num-
likc Dicrochilc; piccous black, lateral mar-
gins ol proiiotiiin ;iiul elytra slightK trans-
The Carabid Beetles of New Guinea • Darlinfiton
17
lucent, elytra iridescent; microsculpture
fine and isodiametric on front, indis-
tinct (at 100 X ) but probably strongly
transverse on pronotum and elytra. Head
0.74 and 0.72 width prothorax; eyes large;
front slightly convex, weakly impressed at
sides anteriorly. Frothorax quadrate-sub-
cordate; width length 1.35 and 1.39; base/
apex 1.13 and 1.09 (base measured across
posterior-lateral setae); base slightly emar-
ginate, not margined; apex broadly emar-
ginate, with impressed marginal line; sides
rounded except c. straight toward base;
margins rather broad, moderately explanate,
each with usual 2 setae ( at base and before
middle); basal angles very obtuse, almost
rounded; pronotum with usual impressions,
impunctate at middle, closely punctate at
base and sides. Elytra subparallel, slightly
narrowed toward base; width elytra/pro-
thorax 1.46 and 1.40; outer-apical and
sutural angles each with an acute tooth or
very short spine; striae shallow, faintly
punctulate; intervals c. flat, subequal on
disc, 3rd with 2 punctures attached to 2nd
stria. Legs: middle and hind tarsi broadly
grooved each side above; 5th segment hind
tarsi with e. 6 strong setae each side below.
Secondary sexual charaeters: i front tarsi
somewhat obliquely dilated, with 3 seg-
ments squamulose below; 6 with 1, 9 with
2 setae before apex each side last ventral
segment. Measurements: length c. 12.5-
15.5; width c. 5.0-6.4 mm.
Types. Holotype c^ (A.M.N.H.) and 1 9
paratype (M.C.Z., Type No. 31,347) from
Lake Daviumbu, Fly R., Papua, Sept. 1-10
(holotype) and Aug. 19-30 (paratype),
1936 (Archbold Exp.), evidently taken in
a light trap; 1 9 paratype (Bishop Mus.),
Oriomo R., Papua, 6 m, Feb. 13, 1964,
"H. C", in light trap; 1 9 paratype, "Highl.
Agr. Exp. Sta./Aiyura, E. Highl./D", N-E.
N. G., 5600 ft. (c. 1700 m), May 26, 1960
(J. J. H. Szent-Ivany, Dept. Agr. Port
Moresby), at light; 1 S paratype (Bishop
Mus.), Nabire, S. Geelvink Bay, West
N. G., 10-40 m, Oct. 7, 1962 (H. Holtmann),
m light trap in jungle.
Measured specimens. The 6 holotype
and 9 paratype from Lake Daviumbu.
Notes. This species is closely comparable
only with D. gigas Castelnau, of Australia.
It resembles gigas in most technical char-
acters including the denticulate-spinose
elytra, but differs from gigas in being much
smaller (Australian gigas measure 20 mm
and over) and in having a relatively nar-
rower prothorax and less impressed front.
Louwerens (Treubia 24, 1958, p. 250)
records D. gigas from Obi Is. in the Moluc-
cas, on the basis of 2 specimens 18 mm long,
which differ in some details from the single
Australian specimen of gigas with which
they were compared. Whether the Obi Is.
specimens are gigas or a related species
remains to be decided, as Louwerens hints.
Dicrochile alternans n. sp.
Description. Form (Fig. 7) of rather
broad Australian Dicrochile (e.g., goryi
Boisduval); black, appendages blackish ex-
cept outer segments of antennae brown;
both sexes moderately shining but not
iridescent, with reticulate microsculpture
faint and c. isodiametric ( where detectable)
on front, vague or irregular but apparently
transverse on pronotum and elytra. Head
0.74 and 0.73 width prothorax; eyes mod-
erate; front convex at middle, irregularly
longitudinally impressed each side anteri-
orly. Frothorax slightly transverse, width/
length 1.36 and 1.38; base/apex 1.15 and
1.15; base and apex broadly emarginate,
apex strongly and base less strongly or in-
distinctly margined; sides broadly rounded;
margins broadly flattened and moderately
reflexed posteriorly, each with usual 2
setae, at base and before middle; basal
angles broadly rounded; disc convex, with
usual impressions, punctate at base, sides,
and apex, impunctate at middle. Elytra
elongate-subquadrate; width elytra/pro-
thorax 1.48 and 1.54; apices sinuate but not
denticulate; striae deep, punctulate; in-
tervals convex, unequal on disc (3rd, 5th,
7th nearly 2x as wide as others at % of
elytral length), 3rd usually 2-punctate with
18 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
punctures near or behind ^.-i and 7;5 of
elytral length, but anterior puncture some-
times duphcated on one or both elytra.
Legs: middle and hind tarsi sulcate each
side above; 5th segments hind tarsi with c.
6 strong setae each side below. Sccondanj
sexual characters: 6 front tarsi dilated and
squamulose as usual in genus; 6 with
1, $ 2 setae each side last ventral segment.
Measurements: length c. 11.5-12.5; width
4.5-5.0 mm.
Types. Holotype $ (M.C.Z., Type No.
31,34(S) and 12 paratvpes from Chimbu Vv.,
Bismarck Rge., N-E'. N. G., 5000-7500 ft.
(c. 1500-2300 m), Oct. 1944 (Darlington);
1 paratype, Feramin, N-E. N. G., 1200-1500
m, June 15-18, 1959 (W. W. Brandt, Bishop
Mus. ); 1 paratvpe, Minj, W. Highlands,
N-E. N. G., 5200 ft. (c. 1600 m), May 20,
1960 (J. 11. Barrett, Dept. Agr. Port
Moresby), by mercury vapor lamp.
Additional material. Papua: 1 c5 , S.
Highlands, Aiyuro nr. Mendi, 1530 m, Oct.
7, "l958 (Gressitt), in light trap. West
N. G.: 16, Wissel Lakes, Urapura-Itouda,
Kamo Vy., 1500 m, Aug. 12, 1955 (Gressitt).
Measured specimens. The i holotype
and 1 9 paratype from Ghimbu Vy.
Notes. The usually 2-punctate 3rd elytral
intervals and the deep, punctulate striae
suggest that this new species is allied to
the common Australian Dicrochilc goryi
Boisduval, but the elytral intervals of goryi
do not alternate in width, and there arc
other smaller differences.
I found th(> Ghimbu specimens under
cover on the ground in fairly open places.
Dicrochile tiro n. sp.
Descriptio)\. Similar to the preceding
(aUerruitis) but larger, with flatter front
and relatively wider prothorax. Head 0.71
and 0.69 width prothorax, lormed as in
alternans except Hatter anteriorly. Vro-
thorax: width length 1.39 and 1.45; base
apex 1.16 and 1.16; otherwise as in alternans.
Elytra: width clytra/prothorax 1.47 and
1.40; most details including alternation of
elvtral intervals c. as in (illcr)unis: 3rd in-
terval 2- to 4-punctate, the number of punc-
tures often different on the 2 elytra of 1
individual (actual punctures on the left
and right elytra of 6 individuals are 2-3,
2-2, 4-2, 2-2, 2-2, 2-3). Measurements:
length 13.5-14.5; width 5.5-6.1 mm.
Types. Holotype 9 (Leiden Mus.) and
7 paratypes (some in M.G.Z., Type No.
31,349) all from Wissel Lakes, West N. G.,
as follows: holotype and 4 paratypes. Lake
Paniai, 1750 m, and 1 paratype, Arabu
Camp, ISOO m, various dates in Sept., Oct.,
Nov. 1939 (H. Boschma, Leiden Mus.); 2
paratypes, Enarotadi, 1800-1900 m, July 31,
Aug. 9, 1962 (Sedlacek).
Measured specimens. One 6 paratype
from Lake Paniai and the 9 holotype, in
this order.
Notes. Sufficiently compared with al-
ternans in the preceding Description and in
the Key to S})ecies of Dicrochile of New
Guinea. Whether tiro is a separate species
or a local form of alternans is uncertain.
The matter is complicated by the occur-
rence of a specimen of alternans in the
Wissel Lakes area.
Genus MICROFERONIA Blackburn
Blackburn 1(S90, Proc. l-iiuicaii Sor. New Sdutli
Wales (2) 4, p. 738.
Sloane 1898, Proc. Linncaii Soc. Xcw South Wales
23, pp. 490-491 (Australian species).
Csiki 1931, Coleop. Cat., Caiabidae, Harpalinae
5, p. 920 (see lor additional rcferenc(^s ).
Ck'nyccnis Andrewes 1933, Trenliia 14. \i. 277
( new s\non\ luy).
Diagnosis. See Key to (Uiura of Licinini.
Description. None re(inirc'd here; see
Notes, below.
Ty})e s))ccies. Oi M icrofcronicL M. adc-
laidae Blackburn, Australia; of Genycerus,
Cx. hicanoides Andrewes, of Ja\a.
Generic distribution. Aii.«^tralia, N<mv
(Fiiiiioa. Ja>a. and presumably inlciAcniiig
islands.
Notes. W'lu'n Anchx'wcs described Gouj-
cerus, he thought the mandiblc\s uniciue
among Licinini, but he was not lamiliar
with the Australian members ol the tribe. 1
l)a\(' seen the t\ pe ol (U'nyccnis lucanoidcs
The Carabid Beetles of New Guinea • Darlington
19
and have a photograph of it, and it seems
to me that the mandibles are eomparable
to those of Microfcronia. The diseovery of
another comparable species in New Guinea
links the Australian and Javan forms geo-
graphically. I therefore tentatively suggest
the synonymy cited above.
Microferonia baro n. sp.
Description. Form as in Figure 8, elon-
gate-oval with very small head; brownish
piceous, legs and antennae slightly reddish;
moderately shining, reticulate microsculp-
ture c. isodiametric on front, transverse on
pronotum, more transverse on elytra. Head
0.51 width prothorax; eyes large, genae
short; 2 setae over each eye; antennae with
3 basal segments glabrous; right mandible
2-dentate, with inner tooth strong and
acute ( left mandible probably c. similar but
partly hidden ) ; front almost evenly convex
except with slight frontal impressions an-
teriorly; clypeus subtruncate, with narrow
transverse membrane; labrum emarginate to
c. middle of length, with lobes equal;
mentum without tooth; ligula and paraglos-
sae apparently subequal, ligula apparently
2-setose; palpi slender except apical seg-
ments of both pairs slightly thickened.
Prothorax: width/length 1.40; base/apex
1.70; base truncate-emarginate, vaguely mar-
gined at middle; apex broadly emarginate,
with marginal line entire; sides rounded
anteriorly, nearly straight toward base, nar-
rowly margined, each with 2 setae, at base
and before middle; disc broadly convex
except depressed baso-laterally, impunctate,
with middle line distinct but transverse im-
pressions c. obsolete. Elytra long-oval;
width elytra prothorax c. 1.30; margins
entire at base, bluntly (almost rectangularly)
angulate at humeri, not distinctly sinuate
toward apex; sutural angles narrowly
rounded; striae fine, irregular but scarcely
punctulate; intervals nearly flat, somewhat
irregular but scarcely alternating; each 3rd
interval with a conspicuous seta-bearing
puncture about % from base, a less con-
spicuous puncture without seta near or be-
hind middle, apparently no more-posterior
puncture. Inner icin(j,s evidently atrophied.
Loner surface almost impunctate but ex-
tensively alutaceous, not pubescent; met-
episterna less than Vj longer than wide.
Legs: tarsi slender, not sulcate above; 5th
segments hind tarsi with 5 long setae each
side below. Secondary sexual characters:
$ with 3 segments each front tarsus mod-
erately dilated, squamulose below; i with
1 seta each side last ventral segment; i
copulatory organs as in Figure 170; ? un-
known. Measurements: length c. 8; width
3.4 mm.
Type. Holotype $ (M.C.Z., Type No.
31,350) from Mt. Wilhelm, Bismarck Rge.,
N-E. N. G., 7000-10,000 ft. (2135-3050 m),
Oct. 1944 ( Darlington ) ; the type is unique.
It was taken on the ground under cover
in mountain rain forest.
Notes. Microferonia baro is more oval
and smaller-headed than M. (Genycerus)
lucanoides (Andrewes) of Java. I do not
have a specimen of lucanoides, and I do
not want to dissect the mouth parts of the
single type of haro (which should be re-
served for specialists in Licinini), but so
far as I can determine the two species are
similar in generic characters although dif-
ferent in detail. ^L baro is larger, more
oval, and smaller-headed than anv Aus-
tralian Microferonia known to me.
(Tribe AMBLYSTOMINI)
(Genus AMBLYSTOMUS Erichson)
Erichson 1837, Kiifer Mark Brandenburg 1, 1, p.
59.
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae 5,
p. 922 ( see for synonymy and additional ref-
erences ) .
Andrewes 1939, Ann. Mag. Nat. Hist. (11) 3, p.
130.
Diagnosis. Small Carabidae with most
technical characters of large-headed Har-
palini but with labrum usually unsymmetri-
cally emarginate and scutellar striae in first
(not second) intervals; length usually less
than 5 mm.
Description. None required here.
20 BuUetin Museum of Comparative Zoology, Vol. 137, No. 1
Type species. Acupalpus mauritaniciis
Dejean, of the Mediterranean region (An-
drewesl939).
Generic distribution. Most of the wanner
parts of the Old World, inckiding Aus-
tralia but perhaps not New Guinea.
Notes. In the British Museum are 14
specimens labeled as from Dor(e)y, New
Guinea, some marked as collected by Wal-
lace and all probably from his material.
They include 4-macuIate, 2-maculate and
immaculate individuals, probabh represent-
ing different species. However, these speci-
mens may be mislabeled and may really be
from Celebes or the Moluccas (see Part I
of my "Carabid Beetles of New Guinea,"
p. 331 ). I have received no other specimens
from New Guinea and found none there
myself, although I collected series of the
genus in the Philippines, so that my collect-
ing methods are evidently adequate to ob-
tain it, and Amhhjstomus is usually common
where it occurs at all. I therefore doubt
its occurrence in New Guinea. I list the
genus here, in parentheses, but see no rea-
son to name or discuss the "Dor(e)y"
species individually.
Tribe CHLAENIINI
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae
5, \). 927 (see for earlier references and
synoinmy ).
Callistitac Jeannel 1942, Faune de France, Coleop.
Carabiques, Part 2, p. 961.
Jeannel 1949, Coleop. Carabifjues de la Rei^ion
Malgache, Part 3, p. 776.
Calllstinae Basilewsky 1953, Exploration Pare Na-
tional I'Upemba, Fasc. 10, Carabitlae. p. 119.
A single, well known genus of this tribe
is represented in New (Guinea.
Genus CHLAENIUS Bonelli
iioiiclii bSlO, Observations Eiit. 1, Tab. Syiiopt..
Mem. .Acad. Sci. Tnrin 18, pp. 21-7(S.
Chaudoir 1876, Ann. Mus. Civ. Genoa 8, p. 10
(in m()n()L,'rai)li of "Chleniens").
Sloane 1910, Proc. Linnean Soc. New South \\ ales
35, p. 437 (Anstralian species).
Csiki 1931, Coleop. Cat., Carabidae, Haipalinae 5,
p. 934 (see for additional references).
Andrewes 1941, Ann. Maji. Nat. Hist. (11) 7, p.
307 (with key to Javan species),
jeannel 1942; 1949 (see works cited nnder tribe).
Bell 1960, Misc. Pub. Ent. Soc. America 1, pp.
98, 108 (North American species).
Diasi,nosis. See works cited. I use
Chlaenius in a xery broad sense, as noted
below. In this sense it is the only genus of
the tribe in New Guinea and Australia.
Description. None required here. For
discussion of some characters of the New
Guinean species, see Notes below.
Tyj)c species. Chlaenius nmrii,inatus Rossi
{= vclutinus Duftschmidt), of Em-ope.
Generic distribution. Nearly world-wide.
Tlie genus is most diverse in structure and
most numerous in species in Africa and the
Oriental Region, less diverse and less nu-
merous in Eurasia and America north of the
tropics, and still less in South America and
the Australian Region. This suggests that
the genus has evolved primarily in the
Old World tropics and spread from there.
In the Asiatic-Australian area, scores of
species of CJda-cnius are known in tropical
Asia, about 30 in Java (Andrewes 1941),
but only 12 in New Guinea, and only 10
(including Ilololeius) in Australia. Some of
the species in New Guinea and Australia
are undifferentiated Asiatic forms. Others
are endemic to New Guinea or Australia.
And the endemics difft^r in degree of dis-
tinctness. This suggests continual trickling
of species from Asia toward New Guinea
and Australia rather than concerted move-
ments. The fact that all Chlaenius in New
Guinea and Australia are still winged sug-
gests that their dispersals haxc been rela-
tively recent.
Notes. Vov discussion ol the authoi-, date j
of publication, and type species of Chhicnius !|
see Jeannel 1942, page 963, footnote.
Chlaenius is a huge genus of 700 or SOO
or more known species, and the species are
diverse and can be di\ided into man\- well
characterized groups. Ne\-ertheless, the
genus as a whole seems natural, not poK-
phyletic. Under these circumstances, al-
though the genus can and should be
subdivided, the taxononiie lexcl of tlu- sub-
dixisions should be detcMinined b\- utilit)'
and intelligibilit) . Chlaenius is known to
The Carabid Beetles of New Guinea • Darlington 21
many entomologists who are not specialists species. Segment 3 is not strictly glabrous
in Carabidae, and there seems much to lose in any species; a few minute setules are
and little to gain by splitting it into small aWays visible in fresh, clean specimens at
genera with new and unfamiliar generic 50x or lOOx magnification.
names, many of them unfamiliar even to The palpi (lioth pairs, in both sexes) are
me, a specialist in Carabidae! I shall there- usually slender with apices narrowly trun-
fore use ChJacnius in a very broad sense, cate, but are almost acuminate in guttula,
as a matter of considered policy. I expect and more broadly truncate in flaviguttatus
to discuss this policy in more detail in Part (tenninal segments with apical edges Vs or
IV of my "Carabid Beetles of New Guinea." % segments' length ) .
The ChJocniiis of New Guinea are few and The mentum is toothed, and the tooth is
some of them do not fit well in recognized usually variably emarginate.
subgenera, and no one is likely to be misled The pronotum has a basal hair fringe in
if I treat them simply as species of the all species except daer, ceylanicus, and
great genus Chloenius sensu Into. guttula, which lack it. These 3 species are
Almost every author who has worked apparently not related to each other,
extensively on C/?/rtcn/n.5 has used new char- The pronotum always has a pair of pos-
acters to group the species, but the works terior-lateral and in some cases also median-
of different authors have not been well cor- lateral seta-bearing punctures, but they are
related. Chaudoir used a variety of obvious often hard to distinguish in the general
characters, beginning with extent of ab- punctation. The posterior-lateral punctures
dominal punctation. Sloane noted that are Vo or Ve of the prothoracic length before
presence or absence of a basal pronotal the base in Chlaenius pan, daer, and guttula,
hair fringe and presence or absence of inter- but closer to or at the posterior angles in
ruptions of the outer elytral margins are the other species. Median-lateral punctures
promising taxonomic characters within the (just before middle of prothoracic length)
genus. Jeannel and Basilewsky derived new are present in some (all?) individuals of
group characters from the male genitalia, occidtiis and siccus, but apparently absent
And Bell found additional characters in the in the other species.
labial pit organs of both sexes and in the The elytra have the basal margin entire
chaetotaxy of the valvulae of the female, except in Chlaenius pan. The margin is
The following notes on certain characters obtusely angulate at humeri in daer,
apply only to the New Guinean species of rounded or at most vaguely subangulate in
Chlaenius, unless otherwise indicated. the other species.
Tlie mandibles are short in all Chlaenius The outer elytral margins are interrupted
in New Guinea, and are exceptionally before apex except in Chlaenius pan and
strongly semicircularly arcuate in maculiger. daer, in which the interruption is obsolete.
The clypeus and labrum are truncate or The punctation of the elytral intervals
weakly emarginate except in amplipcnnis, is 2-seriate in Chlaenius pan and daer, but
in which the labrum is deeply emarginate. irregular in the other species, in which it
The antennae have segment 3 much (c. varies from sparse {ceylanicus only) to
V2) longer than segment 4 in Chlaenius pan dense.
and daer, slightly longer in guttula and The inner wings are full and probably fit
amplipennis, and subequal in the other for flight in all Chlaenius in New Guinea
species. Segment 3 is pubescent in guttula and also in Australia, although wang atrophy
(although the pubescence differs in quality has occurred in various African, Asiatic,
from that on the outer segments), more and North American stocks of the genus,
sparsely pubescent or setulose in pan and The punctation of the lower surface of
daer, and still more sparsely so in the other the body is more diverse than some authors
22 BiiUetin Museum of Coinparative Zoology, Vol. 137, No. 1
have realized. Almost the whole lower
surface including the abdomen is punctate
or punctulate and setulose in CJilaenitis
guttula and amplipennis and also in dacr,
although the latter belongs to the circiiin-
datiis group in which Andrewes (1941)
considered the middle of the abdomen
glabrous. C. pan, ceylanicus, and muculiiicr
are more or less intermediate in this char-
acter. The other species have the middle
of the abdomen widely glabrous.
The metepisterna are differently mar-
gined in different Chlocnhis in New Guinea,
but I doubt if this character deserves the
importance Andrewes (1941) gives it.
The tarsi are obviously setulose above
in CJihicnius U,uttula, glabrous or nearly so
in the other species, but minute setules are
usually visible on the upper surface of the
tarsi at 50 X or l(X)x magnification, even in
the "glabrous" species.
The hind tarsi have the 5th segments
always with 2 rows of strong setae below.
The number of setae in each row varies
from about 4 to about 8 in different species.
Males of all New Guinean species of
CJihicnius have each front tarsus with 3
segments dilated (least so in ccylaniciis,
see following Key ) and densely squamulose
below. And i 6 have 1, 9 9 2 setae each
side before apex of last ventral segment,
with extra adventitious setae som(>times
present.
The aedeagus is open above for much
ol its length in most species (especially
widely open in iiuttiila) but relatively long
and closed lor almost half its length in pcni.
I have not studied the chaetotaxy of the
9 valvulae.
Six unrelated species of CJihicnius, de-
rived from groups that normally have pale
markings on the elytra, are losing or have
lost the markings in New Guinea (see Notes
under (hier, guttula, flaviguttafus, himac-
uhiius jiongraczi, maculiger, and iunnifer
malclicri). This suggests a local climatic
or other selective factor favoring dark color
and loss of markings in New Guinea.
In habits, all New Guinean Chhienius
are ground-living. C. daer, ceylanicus,
himaculatus pongraczi, and occultus are
found on river banks; occultus especially
may occur only beside rivers. C. hamifer
malclicri and siccus are commonly found
under cover in comparatively dry places.
C. maculiger is, I think, a rain forest species.
The other species live in more or less damp
places, but I cannot give their habitats
exactly. I took specimens of several species
at light or in floods.
Key to Species of Chlaenius of New Guinea
1. Elytra with outer margins not interrupted;
elytra! intervals each with 1 re^iular row
of punctures on each side; antennae with
3rd segments c. ^-j longer than 4th 2
- Elytra with outer margins interrupted be-
fore apex; elytral intervals irregularly punc-
tate; antennae witli 3rd segments not or
not much longer than 4th 3
2. Very large (c. 25 nun); pronotmn with
basal hair fringe (p. 23) )Hin
- Smaller (c. 12-15 mm); pronotum without
basal hair fringe (p. 24 ) .— dacr
3. Elytral intervals very sparsely punctulate;
c^ front tarsi narrower, with 2nd segments
V-t or V-i longer tlian wide (p. 24)
ceylanicus
- Elytral intervals more closely pimctulate;
S front tarsi wider, 2nd segment c. wide
as long - 4
4. Mandibles very short, semicircularly arcuate
(p. 25) macidi^cr
- Mandibles normal, moderately arcuate ^^^ 5
5. Abdomen plainK' punctulate and pubescent
or setulose at nu'ddlc as well as at sides 6
- Abdomen broadh' smooth and glaliious
(or nearly so) at middle 7
6. Labrum subtruncate or weakly emarginate;
pronotum without basal liair iringe; pos-
teiior-lat(Mal selae c. la of i^rothoracic
length before base; size very small ( c.
8 mm) (p. 25) •guttula
- Labniiii deeply emarginate; pronotum with
basal hair Iringe; posterior-lateral setae
near ( slighlK rounded ) posterior angk'S;
larger ( r. 12 nmi ) {p. 26) --- (iiniiliiunnis
7. Male bout leniora each with a small tooth-
like tub('rel<' Ix'low, near base; 5th seg-
mcnls liiiul tarsi with c. 7 or cS setae each
side below S
- Male front b^nora without tubercles; 5th
segments hind taisi with r. 5 setae each
side below 10
The Carabid Beetles of New Guinea • Darlington 23
8. Pionotum closely and coarsely punctate
and head including front punctulate ( this
combination of characters separates both
sexes of this species from all following
ones, the closest approach being siccus,
see couplet 10) (p. 26) flaviguttatus
— Pronotum with only base coarsely punctate;
head not or only sparsely irregularly
punctulate 9
9. Pronotum with anterior margin entire or
only narrowly interrupted at middle; sides
of prothorax usually sinuate; elytra usually
2-niaculate ( p. 27 ) -.- hiimiculatus pongraczi
— Pronotum with anterior margin obsolete,
indicated only toward sides; sides of pro-
thorax not sinuate; elytra not maculate
( p. 27 ) olthofi
10. Pronotum punctate at base and in narrow
zone along midline but much of disc im-
punctate; posterior-lateral pronotal setae
often c. 1/4(1 of pronotal length before angles
(but variable) (p. 28) occultus
- Pronotum more extensively ( but not always
evenly) punctate; posterior-lateral pronotal
setae almost at basal angles 11
11. Pronotum more sparsely punctate near mid-
dle, with punctiues tending to form ir-
regular longitudinal rows; front exten-
sively but irregularly punctate (p. 28) _^_-
Jimnifcr malcheri
- Pronotum coarsely pimctate, with punc-
tm^es somewhat irregular but less so than in
preceding species; front shining, widely
impunctate ( a few punctines posteriorly
and laterally); (this species characterized
also by coarse punctation of proepisterna
and of elytral striae) (p. 29) siccus
Chlaenius pan n. sp.
Description. Form as in Figure 9, large,
rather slender; black, appendages brownish
piceoiis except c. outer halves of femora
reddish testaceous; rather shining; reticulate
microsculpture fine, faint on front, slightly
more distinct on pronotum and elytra, c.
isodiametric except slightly transverse on
part of pronotum. Head 0.81 and 0.84 width
prothorax; eyes rather abruptly prominent;
antennae with 3rd segments about V.>
longer than 4th segments and plainly but
sparsely setulose; mandibles short, mod-
erately arcuate; mentum with deeply emar-
ginate tooth; clypeus subtruncate; labrum
slightly emarginate; palpi narrowly truncate
at apex in both sexes. Fvothorax quadrate;
width/length 1.11 and 1.09; base/apex 1.16
and 1.11; sides weakly arcuate anteriorly,
slightly converging and very broadly weakly
sinuate posteriorly, each with seta c. Vt> of
length before base, without median-lateral
seta; disc with impressed middle line and
rounded basal impressions, wrinkled-punc-
tate at base, nearly smooth (sparsely punc-
tulate) elsewhere; posterior pronotal hair
fringe present. Elytra long, narrowed to-
ward base; width elytra/prothorax 1.67 and
1.75; margins c. obliterated at base (inside
bases of 4th striae), rounded at humeri, not
interrupted posteriorly; intervals rounded-
subcostate, each with an irregular row of
punctures on each side. Lower surface
partly irregularly punctulate, but much of
abdomen smooth at middle; metepisterna
long, with outer edges raised but not
channeled. Inner wings full. Legs slender;
tarsi not pubescent above; 5th segments
hind tarsi with 4 or 5 strong setae each side
below. Secondary sexual characters normal;
2nd segments S front tarsi c. wide as long
(by measurement); d front femur not
dentate; aedeagus long, slender, closed
above for nearly half its length (Fig. 171).
Measurements: length c. 25-26; width c.
9.1 mm.
Types. Holotype $ (Bishop Mus. ) from
Torricelli Mts., Mokai Village, N-E. N. G.,
750 m, Jan. 1-23, 1959 (W. W. Brandt);
and paratypes as follows. N-E. N. G.: 1
$ , Maprik, Sepik Dist., 19^5 ( Dept. Agr.
Port Moresby). West N. G.: 1 9 (M.C.Z.,
Type No. 31,351), Kota Nika, Res. Hol-
landia, Jan. 9, 1958 (R. T. Simon Thomas);
1 9 , Tanahmerah, Res. Boven Digoel, Feb.
1958 (R.T.Simon Thomas).
Measured specimens. The S holotype and
9 paratype from Kota Nika.
Notes. This new species probably repre-
sents Chlaenius femoratus Dejean of Java,
Sumatra, etc. but is narrower (especially
the jjrothorax) and duller than femoratus
and lacks subapical interruptions of the
elytral margins, which are present though
weak in my 5 specimens of femoratus from
Java. I have 1 9 of a related undescribed
s^Decies from Celebes, which partly fills the
24 Bulletin Museum of Comparative Zoologij, Vol. 137, No. 1
geographic gap between femoratus and
pan.
Chloenius doer n. sp.
Description. Form of Chlacnius of cir-
cumdatus group; slender; greenish l)lack,
head green, elytra sometimes with vestige
of very narrow yellowish margin at apex,
appendages testaceous brown; reticulate
microsculpture alisent or faint on head and
pronotum, deep, fine, isodiametric on
elytra. Head 0.82 and 0.(S2 width prothorax;
eyes large, prominent; antennae with 3rd
segments c. ^2 longer than 4th, setulose;
mandibles moderate; clypeus and labrum
subtruncate; surface of head irregularly, not
densely punctate; mentum with ± emargi-
nate tooth. Prothorax narrow, quadrate-sub-
cordate; width length 1.11 and 1.16; base/
apex 1.04 and 1.09; sides arcuate except
broadly usually strongly sinuate posteriorly;
margins narrow, each with seta c. % of
length before base, without median-lateral
seta; disc irregularly punctate, with fine
middle line, linear baso-lateral impressions
nearer sides than middle but shallower than
usual in the group; posterior pronotal hair
fringe absent. Elytra slightly narrowed an-
teriorly; width elytra/prothorax 1.63 and
1.66; margins entire at base, obtusely angu-
late at humeri, not interrupted pcxsteriorly;
intervals weakly convex, each with a row of
punctures on each side. Lower surface in-
cluding middle of abdomen extensively
punctulate and pubescent; metepisterna
long, w(>akly margined externally. Inner
winii-s full. Lcii^s: tarsi nearly glabrous
above; 5th segments hind tarsi with c.
4 short setae each side below. Secondary
sexual characters normal: 2nd segment male
front tarsi c. Vio longer than wide; male
femora not dentate; aedeagus open abov(>
for much of length. Measuremenls: length
c. 12-15.5; width 4.4-6.0 mm.
Types. Holot>T)e i (M.C.Z., Type No.
31,352) and 2 paratypes from Nad/ab, N-E.
N. G., July 1944 (Darlington); and addi-
tional paratypes as follows. Papua: 1<S,
Kiunga, Fly U., dates in July, Aug. 1957
(W. W. Brandt, Bishop Mus.); 1, Lake
Da\iumbu, Fly R., Aug. 19-30, 1936
(Archbold Exp., A.M.N.H.); 1, Palmer R.
at Black R., July 22-31, 1936 (Archbold
Exp., A.M.N.H.); 1, Kokoda, 1200 ft. (366
m), Aug. 1933 (Cheesman). N-E. N. G.: 2,
Aitape, Aug. 1944 (Darlington); 1, Main R.,
Sepik, Feb. 1965 (R. Hornabrook). West
N. G.: 1, Hollandia, July-Sept. 1944
(Darlington); 1, Tanahmerah, Boven Di-
goel Res., 17 m, April 15, 1955 (L. D.
Brongersma, Leiden Mus.); 1, Idenburg R.,
400 m, July 15-Sept. 15, 193S (J. Olthof,
Leiden Mus.); 1, lebele Camp, Snow Mts.,
2250 m, Sept. 1938 (Toxopeus).
Measured specimens. The i holotype and
1 9 paratype from Kiunga.
Notes. C. daer is the only species of the
Chlaenius circumdatus group in New
Guinea. This group is widely distributed
and common in the warmer part of the Old
World including Australia. The present
new species seems nearest acroxanthus
Chaudoir (which ranges from the south-
eastern corner of Asia to the Moluccas —
Louwerens 1956, Treubia 23, p. 223) but
has baso-lateral pronotal impressions shal-
lower and punctation less coarse. I have
used for comparison a series of acroxantJuis
from Java, collected by Thomas Barbour.
Chlaenius ceylanicus Nietner
Nietner 1856, j. Asiatic Soc. Bengal 25, p. 385.
Csiki 1931, Cok'op. Cat., Carabidat', ilarpalinae
5, p. 932 (HoloUns) (.see for additional refer-
ences ).
niliditlus Dejean (not Sehraiik) 182(i, Species
General Coleop. 2, p. 34 1 .
onuitus Tryon 1890, Second Annual Report
Adnn'nistrator British New (Guinea, Appendix 5,
p. 109 (Poecilus).
Csiki 1931, Coleop. ("at., Carahidae, Ilarpalinae 5,
p. 563 (?Focciloi(lca).
Descri])lion. None recjuired here; see
preceding Key to Sj)ccies for recognition
characters. Note 2nd segments 6 front tarsi
'1 (Ja\;ui specimen) or ':i (Australian speci-
men) longer than wide (l)y measurement);
length c. 11-12.5 mm.
Types. Of ceylanicus. from w(\stern and
southern Cevloii; now in I^erlin U. Zool.
The Carabid Beetles of New Guinea • Darlington 25
Mus. and Stettin Town Mus. (t. Andrewes).
Of nitidulus, from "Indes orientales"; in
Oberthiir Coll., Paris Mus. Of omofus. a
6 from St. Joseph (Angabimga) R. District,
Papua, collected by A. C. English; present
location of type unknown (not seen).
Occurrence in Neic Guinea. Papua: the
type of ornafus; 1, Rouku, Morehead R.,
Apr. 1962 (W. W. Brandt, C.S.I.R.O.).
N-E. N. G.: 5, Kamindibit, Main R., Sepik,
Feb. 1965 (R. Hornabrook), on water
weeds in swamp. West N. G. : 1 9 ,
Garian, Lake Jamoer, Dec. 8, 1954 (L. D.
Brongersma, Leiden Mus.).
Notes. ''Hololius" ceyJonicus ranges from
southern Asia to eastern Australia, and
will probably be found on all the inter-
vening islands, although records are still
incomplete. I have found it in Australia
under cover by backwaters of rivers and in
river floods. Nietner says it flies to light
in Ceylon.
Chlaenius maculiger Casteinau
Castelnau 1867, Notes on Australian Colcop., p. 62.
Chandoir 1876, Ann. Mus. Civ. Genoa 8, p. 67.
Sloane 1910, Proc. Linnean Soc. New South Wales
35, pp. 438, 440.
Csiki 1931, Coleop. Cat., Caraliidae, Harpalinae
5, p. 961.
nigripes Macleay (not Dejean, not Faldermann )
1886, Proc. Linnean Soc. New South Wales
(2) l,p. 140.
biroi Csiki 1931, Coleop. Cat., Carabidae, Har-
palinae 5, p. 948.
Description (for recognition only). Me-
dium sized, depressed; dark, dark-legged,
typically 2-maculate but spots sometimes
lost; unique in genus in New Guinea in
mandibles very short, semicircular; length c.
12-14 mm.
Types. Of maculiger, from Rockhamp-
ton, Australia; probably in Genoa Mus. ( I
did not find it at Melbourne in 1957). Of
nigripes, from Fly R., Papua (implied);
may now be in Macleay Mus., Sydney (not
seen). Of hiroi, as for nigripes (the name
hiroi was proposed to replace nigripes
Macleay, which is preoccupied).
Occurrence in New Guinea. Widely dis-
tributed and common: 127 specimens from
localities including Dobodura and Wau
in all 3 political divisions of New Guinea;
most at low altitudes but reaching at least
1300 and 1500 m at Wau and on the
Bismarck Rge.
Notes. Sloane (1910) has established the
identity of nigripes Macleay (hiroi Csiki)
with maculiger Castelnau.
Outside New Guinea, this species is
known from Australia and New Britain
(Cape Gloucester, Jan.-Feb. 1944, Darling-
ton). It is apparently related to and
probably derived from the same Oriental
stock as Chlaenius tetragonoderus Chaudoir,
which is widely distributed farther west in
the Malay Archipelago, to the mainland
of Asia. C. tetragonoderus batjanicus
Louwerens (1956, Treubia 23, p. 234) of
the northern Moluccas, which varies in
color of legs, may be a transitional form.
An apparently undescribed species of the
group occurs in the Solomons.
The yellow subapical elytral spots are
individually variable in specimens from
New Guinea and are absent or nearly
absent in some individuals. The variation
in spotting apparently occurs throughout
New Guinea.
This is, I think, a rain forest species that
may occur in ordinary leaf litter rather
than in specially wet places, but I have
taken too few specimens to be sure.
Chlaenius gutfula Chaudoir
Chaudoir 1856, Bull. Soc. Nat. Moscow 29, Part
2, p. 216.
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae
5, p. 957 ( see for additional references ) .
Andrewes 1941, Ann. Mag. Nat. Hist. (11) 7, p.
310 (in key).
Louwerens 1953, Verhandlungen Naturforschenden
Gesellschaft Basel 64, p. 313.
csikii Jedlicka 1951, Ann. Mus. National Hungary
1, p. 136 (new synonymy).
astrolabe nsis Jedlicka 1951, Ann. Mus. National
Hungary 1, p. 136 (new synonymy).
immaculata Louwerens 1962, Tijdschrift voor Ent.
105, p. 145 (new synonymy).
Description ( for recognition only ) . Very
small; dull dark bluish, with or without a
small pale spot on suture near apex (see
26
BuUctin Museum of Comparaiive Zoolop^ij, Vol. 137, No. 1
following Notes); see also preceding Key
to Species; length c. 8 mm.
Types. Of gtittiila, from Hongkong; in
Oberthiir Coll., Paris Mus. Of r.s//\// and
astrolahensis, both from Stephansort, Astro-
labe Bay, N-E. N. G.; in Hungarian
National Mus. Of immacuJata, from Ani-
hoina; in Louwerens Coll. (Types not
seen. )
Occurrence in New Guinea. Papua: 7,
Dobodura, Mar.-July 1944 (Darlington);
1, Bisianumu, near Sogeri, 500 m. Mar. 15-
20, 1955 (E. O. Wilson, M.C.Z.), in rain
forest; 1, Brown R., May 23, 1956 (E. J.
Ford, Jr., Bishop Mus.). N-E. N. G.: 1,
Bulolo, 731 m, Aug. 26, 1956 (E. J. Ford,
Jr., Bishop Mus.); 1, Finschhafen (L.
Wagner, M.C.Z.). West N. G.: 3, Hol-
landia, Jan., Apr., Mav 1945 (B. Malkin,
U.S.N.M.); 1, Kota Nika, Res. Hollandia,
Nov. 29, 1957 (R. T. Simon Thomas,
Louwerens Coll.); 1, Maffin Bay, Jan. 1,
1945 (E. S. Ross, California Acad.).
Notes. This species is known from south-
ern Asia, Sumatra, Java, Bali, Celebes,
Timor (Louwerens 1953), the Philippines,
New Guinea, and New Britain (Cape
Gloucester, Darlington, M.C.Z. ). It often
flies to light.
Although most specimens from New
Guinea have a variable ( sometimes minute )
vestige of a subapical sutural pale spot,
several ( not all ) of those from Dobodura
are unspotted.
The characters given by Jedlicka to dis-
tinguish csikii from iiuttuJa seem to me to
be individual rather than spcx'ific, and
"aberration" astrolalmnsis Jedlicka and "var."
inimaculata Louwerens are (I think) un-
necessary names lor imspotted individuals.
Chlaenius amplipennis Chaudoir
Cliaiuloii 1876, Ann. Mus. Civ. Ocnoa 8, p. 252.
Csiki 1931, Coleop. Cat., Carabidac, Ilarpalinac
5, p. 946.
Andrewi's 1911, Ann. Mas. Nat. Hist. (II) 7,
p. 310.
Description (for recognition only). Me-
dium small; dark, dull; uiu(iue among
Chlaenius of New Guinea in labrum deeply
emarginate; length c. 12 mm.
Type. A $ from Java; in Brussels Mus.
(not seen).
Occurrence in New Guinea. N-E. N. G.:
1 6, Bulolo, 2000 ft. (610 m), Mar.-July
1937 (George Rio, Chicago Mus.); 1 9,
Main R., Sepik, Feb. 1965 (R. Hornabrook).
Notes. Chlaenius amplipennis apparently
ranges from Sumatra and Java to the
Philippines, New Guinea, and the Solo-
mons ( Guadalcanal Is., 1944, L. N. Jarcho,
M.C.Z. ). It varies geographically and some
of the geographic fonns may be recogniz-
able subspecies, but I do not have enough
material to decide about this.
Chlaenius flaviguftafus Macleay
Macleay 1825, Annulosa Javanica, p. 14.
Chaiuloir 1876, Ann. Mus. Ci\\ Cenoa 8, p. 52.
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae
5, p. 955 ( see for synonymy and additional
references ) .
Andrewes 1941, Ann. Mati. Nat. Hist. (11) 7,
p. 307 (in key).
Louwerens 1953, Verhandlungen Naturforschenden
Cesellschaft Basel 64, p. 311.
fliittatiis Eschscholtz 1833, Zoologisehen Atlas 5,
p. 26, pi. 25, fig. 8.
iiuniactilipcnuis Jedlieka 1951, Ann. Mus. National
Hungary 1, p. 134 (new synonynn' ).
Description (for recognition onU). Me-
dium sized, rather slender; dull, dark, elytra
2-maculate or immaculate, legs pale usually
with dark knees; most ol upper surface
closely conspicuously punctate; palpi with
apical segments truncate, apices -i or ^-i
wide as lengtli of segment; see also Key to
Species; length c. 11-14.5 mm.
Types. Of flaviL!,uttafus, from Java; in
Briti.sh Mus. (seen). Of iiuttatus. Ma-
nil (l)a; Moscow U. Zool. Mus. (not seen).
Of it)nn(iculi))cnnis. New (hiinea; in Jed-
licka ('oil. (not seen).
Occurrence in Neu Ciuinca. Widely
distributed on Ne\\- Guinea ( including
Doboduia and Wan), and reaching I^iak
and (in tUv .Vdmiralties ) Manns Is.: 283
specimens, most at low altitudes but reach-
ing at least 1300 and 1500 m in places in
the mountains.
The Carabid Beetles of New Guinea • DaiUngton 27
Notes. This common Chlaenius is known
from Sumatra, Java, etc. to New Guinea
and Australia, east to the Philippines,
New Britain, New Ireland, Solomons,
New Hebrides, Fiji, Samoa, and New
Caledonia.
Markings vary indwiduaUij in the series
from New Guinea. Each elytron may have
a conspicuous irregular subapical pale mark,
or fragments of such a mark, or no mark at
all, and the variation occurs in all parts of
New Guinea from which series have been
seen. Unmarked individuals are "aberra-
tion" immacu]ipcnms]ed\\ckA, which I think
is not worth distinguishing.
This species occurs in a variety of wet
places, often in more or less open country.
Chlaenius bimaculafus pongraczi Jedlicka
Jedlicka 1951, Ann. Mus. National Hungary 1,
p. 136.
Description. Generally similar to typical
himaciilatus Dejean in technical characters
( see preceding Key and also Andrewes' key
to Javan Chlaenius, 1941, Ann. Mag. Nat.
Hist. (11) 7, p. 307), but differing some-
what in color and especially in punctation.
Color bluish black, legs testaceous (not
bicolored), antennae and mouthparts red-
dish brown. Punctation of head and pro-
notum reduced but variable: head with or
almost without punctulation (most distinct
posteriorly); pronotum coarsely punctate
only basally, extensively smooth or in part
finely punctulate elsewhere; length c. 12-
14 mm.
Type. From New Guinea; in Hungarian
National Mus. (not seen).
Occurrence in New Guinea. Papua: 9,
Dobodura, Mar.-July 1944 (Darlington);
7, Mt. Lamington, 1300-1500 ft. (c. 400-
450 m), (C. T. McNamara, S. Australian
Mus.); 2, Kokoda, 1200 ft. (366 m). May
& Aug. 1933 (Cheesman); 1, Daradae, near
Javarere, Musgrove R., Oct. 4, 1958
(Gressitt). N-E. N. G.: 3, Sattelberg
(British Mus.); 1, Wareo, Finschhafen
(Rev. L. Wagner, S. Australian Mus.); 1,
Gewak, Salawaket Rge., 1530 m, Sept. 7,
1956 (E. J. Ford, Jr., Bishop Mus.); 1,
Sepik, Maprik area, 160 m, Aug. 26, 1957
(D. Elmo Hardy, Bishop Mus.). West
N. G.: 1, Hollandia, Jan. 1933 (A.M.N.H.);
1, VVaris S. of Hollandia, 450-500 m, July
24-31, 1959 (T. C. Maa, Bishop Mus.); 2,
Ifar, Cyclops Mts., 300-500 m, June 23-25,
Sept. 9, 1962 (Sedlacek); 1, Guega, W. of
Swart Vy., 1200 m, Nov. 14, 1958 (Gressitt);
1, Bodem, 11 km SE. of Oerbefareh, 100 m,
July 7-17, 1959 (T. C. Maa, Bishop Mus.).
Notes. Chlaenius himaculatus Dejean (or
the group of closely related species that
goes under this name) ranges from SE.
Asia to the Philippines and New Guinea
(not Australia). I have ample comparative
material from a number of localities from
SE. Asia to Amboina. Geographic variation
is obvious. The New Guinean form varies
also individually in marking: most individ-
uals have a conspicuous pale spot before
apex of each elytron, but the spot varies
in size and is almost absent in 2 of the
Sattelberg specimens.
My Dobodura specimens were taken in a
grassy bank beside a small river.
Chlaenius olfhofi n. sp.
Description. Form (Fig. 10) of Chlaenius
himaculatus Dejean, slender; head and
pronotum shining green or greenish black;
elytra duller, purplish black, with very fine
c. isodiametric microsculpture; appendages
rufous. Head 0.72 and 0.76 \\'idth prothorax;
eyes large, genae short; antennae with 3rd
segments c. equal to 4th and sparsely
setulose; mandibles average; clypeus and
labrum subtruncate or weakly emarginate;
front with c. punctiform anterior impres-
sions, otherwise almost impunctate (a few
punctules posteriorly); mentum with c.
entire tooth; palpi narrowly truncate at
apex. Prothorax subquadrate, widest at or
slightly behind middle; width/length 1.16
and 1.13; base/apex 1.17 and 1.10; apex not
margined except vaguely at sides; sides
broadly rounded, not or at most faintly
sinuate before obtuse but well defined,
slightly blunted posterior angles; posterior-
28 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
lateral setae c. Vin of prothoracic length be-
fore base, median-lateral setae absent; disc
with impressed middle line, sublinear
baso-lateral impressions; surface extensively
smooth but with a few punctmes mosth'
near base, sides, and along middle; posterior
pronotal hair fringe present. Elytra: width
elytra prothorax 1.44 and 1.45; margins
entire at base, rounded at humeri, with
subapical interruptions; striae moderately
impressed, vaguely punctulate; intervals
slightly convex, moderately closely punctate.
Lower sui'facc shining; proepisterna almost
impunctate; some punctures on metepisterna
including epimera and on base of abdomen
at sides, but much of abdomen smooth or
nearly so; metepisterna long, margined ex-
ternally, margin obsolete anteriorly. Inner
irin^.s full. Lci^s: tarsi c. glabrous above;
5th segments hind tarsi with c. 7 setae each
side below. Secondary .sexual characters:
i front tarsi with 2nd segments c. wide as
long; <^ front tibiae with small tooth below
near base; c^ with usually 1, 9 2 or 3
( unsymmetric in the single 9 ) setae each
sidc> last ventral segment. Aedeagus slender,
open above for much of length. Mea.siire-
ments: length c. 13-14; width c. 4.5-5.0
mm.
Types. Holotype $ (Leiden Mus.) and
i i paratype (M.C.Z., Type No. 31,353)
from Bcrrihard Camp, West N. (i., 50 m,
July-Sept. 1938 (J. Olthof); 1 9 paratype,
same locality, Apr. 12, 1939 (Toxopeus);
1 i paratype, Oro Bay, Pa|»ua, JuK' 12,
1944 (A. II. Mallery, Bishop Mus.). '
Measured specimens. The -' holotype and
9 paratype.
Notes. This seems to be a distinct species
of the himaculatus group, occurring within
the geographic range of ])iinaculalus sub-
species pon^raczi. C. olthof i may be a prod-
uct of an early invasion of a hinuuulatus-
like stock, pon<^raczi of a later one.
Chlaenius occulfus Sloane
Sloane 1907, Dciitsclie Kiit. Zcitscln ill (ni 1!)()7.
p. 467.
Descrij)lion (for recognition onlv). A
medium-sized Chlaenius with subcordate
prothorax; blue-black, sometimes in part
greenish, legs reddish testaceous, antennae
and mouthparts reddish brown; rather
shining, reticulate microsculpture absent or
nearly so on head and pronotum, visible on
elytra especially of female, fine, irregularly
isodiametric; see also Key to Species of
Chlaenius of New Guinea; length c. 14-
17 mm.
Type. From Herbertshohe, New Britain,
"returned to Dr. Horn for Bennigsen's col-
lection" (not seen).
Occurrence in New Guinea. Papua: 7,
Dobodura, Mar.-July 1944 (Darlington);
4, Kokoda, 1200 ft. (366 m), Mav, Aug.
1933 (Cheesman); 1, Laloki, 1909 (F. Muir,
Hawaiian Sugar Planters Association ) ; 5,
Mt. Lamington, 1300-1500 ft. {c. 400-150
m) (C. T. McNamara, S. Australian Mus.);
1, Peria Ck., Kwagira R., 50 m, "No. 7,"
Aug. 14-Sept. 6, 1953 (Geoffrev M. Tate,
A.M.N.H.). N-E. N. G.: 30, vie. Nadzab,
Julv 1944 (Darlington); 1, Busu R., E. of
Lae, 100 m, Sept.'" 14, 1955 (Gressitt); 2,
Wan, 1100, 1200 m, Oct. 30, 1961, Jub-
28-29, 1963 (Sedlacek). West N. G.: 5.
Hollandia, Jan., Apr., Mav 1945 ( B. Malkin,
U.S.N.M.); 1, Humboldt Bay Dist., 1937
(W. Stiiber, British Mus.); 1, Tanahmerah.
Res. Boven Digoel, Apr. 24, 1957 (R. T.
Simon Thomas, Leiden Mus.).
Notes. I have identified this species from
Sloanes description: the size, cordate pro-
thorax, and round(xl humeral margins are
(together) diagnostic in th(^ region in (pies-
tion; other dc^tails agree well enough; and
I have seen a specimen from New Britain
(near Habaul, l''eb. 1929, Pemberton col-
lector, in Goll. Ilawaiiaii Sugar Planters
Association).
This species occurs in New Britain ( [hv
t\iK\ and tlu> specimen Iroin near iiabaul
reierred to abox'c ) and the Soloiiioim
(Guadalcanal; Bougainville) as wtll as
eastern and central New (Guinea. I ha\e
been unable to determine its relationship to
other si)ecies ol (Chlaenius. It is h)un(l
under stones on the banks ol lixcrs.
The Carabid Beetles of New Guinea • Darlington
29
Chlaenius hamifer malcheri Van Emden
\'an Emden 1937, Stettiner Ent. Zeitung 98, pp.
35, 37.
Description (for recognition only). Me-
dium small, moderately broad; usually
very dark with or without slight metallic
tinge, usually without spots but latter some-
times partly developed (see Notes below);
see also Key to Species; length c. 11-12 mm.
Type. From Pauru, New Georgia, Solo-
mon Islands (Fr. Malcher); in Van Emden
Coll., British Mus. (seen).
Occurrence in New Guinea. Papua: 13,
Dobodura, Mar.-July 1944 (Darlington);
1, Oro Bay, July 12, 1944 (A. H. Mallery,
Bishop Mus.); 1, Kokoda, 1200 ft. (366 m),
Aug. 1933 (Cheesman); 1, Port Moresby,
Konedobu, Apr. 20, 1958 (J. J. H. Szent-
Ivany, Dept. Agr. Port Moresby), at light;
1, Popondetta, Aug. 11, 1962 (A. Catley,
Dept. Agr. Port Moresby), at light; 1, Mt.
Lamington, 1300-1500 ft. (c. 400-450 m)
(C. T. McNamara, S. Australian Mus.); 2,
Rouku, Morehead R., Apr. 1962 (W. W.
Brandt, C.S.I.R.O.); 1, Rossel Is. (S.E.
Papua), Oct. 1963 (W. W. Brandt, C.S.I.
R.O.). N-E. N. G.: 2, "No. 2, Oomsis," 22
mi. W. of Lae on Lae-Bulolo Road, 100 m,
Apr. 26, 1959 (L. J. Brass, A.M.N.H.); 2,
Wau, Morobe Dist., 1200 m, Dec. 6, 1961,
Sept. 15-30, 1962 (Sedlacek); 1, Bulolo "G.
T.," (Sedlaceks); 1, 16 km W. of Mumeng,
3000-5000 m, May 1962 (Sedlacek); 1,
Okapa, July 12, 1964 (R. Hornabrook); 1,
lower Busu R., Huon Pen., May 12, 1955
(E. O. Wilson, M.C.Z.). West N. G.: 1,
Hollandia, May 1945 (B. Malkin, U.S.
N.M.); 1, Maffin Bay, June 1944 (E. S.
Ross, California Acad.); 1, Wissel Lakes,
Tage Lake, 1760 m, Aug. 4, 1955 (Gressitt);
1, Wissel Lakes, Enarotadi, 1900-2000 m,
July 2-11, 1962 (N. Wilson, Bishop Mus.).
Notes. This species belongs to a difficult
group of Chlaenius that extends from S.
Asia to NE. AustraHa. The group includes
hamafus Dejean as well as hamifer Chaudoir.
I am not sure whether these two species
really are different, or with which of them
malcheri should go. My treatment of it as
a geographic form of hamifer is tentative.
The range of hamifer is from S. Asia to
Australia.
Apparently only one form of the hamifer-
hamatus group occurs in New Guinea. It
is very dark and usually unmarked, but 2
examples from Dobodura show the posterior
part of a pale "comma" on the apex of each
elytron, and the 1 specimen from Port
Moresby, the 2 from Rouku, and the 1 from
Hollandia have the "commas" complete but
narrow. Chlaenius insulanus Louwerens
(1956, Treubia, 23, p. 234) of the northern
Moluccas is another dark, unmarked form
of the hamifer-hamatus group, but is smaller
and narrower than malcheri.
C. h. malcheri occurs under cover often
in somewhat drier places than most other
Chlaenius except the following (siccus).
Chlaenius siccus n. sp.
Description. Form c. average in genus;
rather shining black, sometimes with slight
greenish or bluish reflections, appendages
rufous; reticulate microsculpture absent on
head and pronotum, fine and c. isodiametric
on elytra. Head 0.68 and 0.67 width pro-
thorax; eyes large, genae short; antennae
with 3rd segments c. equal 4th and scarcely
setulose; mandibles average, rather short,
moderately arcuate; labrum and clypeus
subtruncate; front almost smooth at middle,
punctate at sides and posteriorly, with
slight frontal impressions; mentum with
blunt usually vaguely emarginate tooth;
palpi slender, narrowly subtruncate at apex.
Prothorax subquadrate but rather strongly
narrowed anteriorly; width length 1.28 and
1.28; base/apex 1.33 and 1.31; sides weakly
arcuate for most of length, c. straight and
somewhat converging posteriorly; posterior
angles obtuse, narrowly rounded; margins
narrow anteriorly, wider posteriorly, each
with posterior-lateral setae just before base
and ( at least in some individuals ) median-
lateral setae just before middle; disc ir-
regularly longitudinally impressed each side
30
Bulletin Museum of Comparative Zoology, Vol. 137. No. 1
c. midway between middle and side, with
whole surface rather closely but somewhat
irregularly, coarsely punctate; posterior pro-
notal hair fringe present. Elytra not nar-
rowed anteriorly; width elytra prothorax
1.29 and 1.36; margins entire at base,
arcuate at humeri, interrupted subapically;
striae rather coarsely punctate, intervals
slightly convex, irregularly punctulate.
Lower swface: proepisterna coarsely punc-
tate at least in part, mesepisterna partly
punctate or almost impunctate, sides of
metasterna punctate, abdomen punctulate
at sides and across base but extensively
smooth or nearly so at middle; metepisterna
long, strongly margined ( grooved ) exter-
nally. Inner uin^s full. Le<!,s without obvi-
ous unusual characters; tarsi c. glabrous
above; 5th segments hind tarsi with a. 4
setae each side below. Secondary sexual
characters normal; i front tarsi dilated, 2nd
segment at least as wide as long; 6 front
tibiae not toothed; c^ with 1, 9 2 setae each
side last ventral segment. Measurements:
length e. 11. .5-13.5; width 4.1-4.8 mm.
Types. Holotype S (M.C.Z., Type No.
31,354) and 14 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. Papua: 2,
Mt. Lamington,' 1300-1500 ft. (c. 400-450
m) (C. T. McNamara, S. Australian Mus.).
N-E. N. G.: 1, Aitape, Aug.-Sept. 1936
(Cheesman); 4, Swart Vv., Karubaka,
1450, 1500, 1550 m, Sept. s', 16, 22, 1958
(Gressitt), some taken in light trap; 8,
Wan. Morobe Dist., 1200 m, dates in Jan.,
Feb., Mar., Aug., 1962-1963 (Sedlacek);
1, Sum-Sum, near Bulolo, Morobe Dist.,
Feb. 7-11, 1966 (Rhonda M. Stevens, Dept.
Agr. Port Moresby); 7, Minj, W. High-
lands, 5200 ft. (c. 1600 m). Mar. 25, May
20, 1960 (J. H. Barrett, Dept. Agr. Port
Moresby). West N. G.: 1, Ilollandia, May
1945 (B. Malkin, U.S.N.M.); 1, Kota Nika,
Res. Ilollandia, Feb. 23, 1956 (R. T. Simon
Thomas, Louwerens Coll.); 4, Ifar, Cyclops
Mts., 300-500 m, June 23-25, 1962 (Sed-
lacek); 1, same locality, 400-800 m, Sept. 9,
1962 (Sedlacek); 2, 'X;. den Iloed. Tfar,"
Dec. 1957 (Louwerens Coll.); 1, Kebar Vy.,
W. of Manokwari, Vogelkop, 550 m, Jan. 4-
31, 1962 (S. & L. Quate, Bishop Mus.).
Measured s))ecimcns. The i holotype and
1 9 paratype from Dobodura.
Notes. In Andrewes' ( 1941, see reference
under genus) key to the Chlaenius of Java,
this runs to leucops Wiedemann, of which
I have specimens from India, Java, and
Luzon, but the present new species is nar-
rower, paler-legged, and more shining than
leucops, with somewhat different sculpture:
e.g., the punctation of the pronotum is
coarser and more irregular than in leucops.
C. siccus is closer to, and may prove to be a
geographic representative of, ophonoides
Fairmaire of Australia ( recorded also from
New Caledonia and New Hebrides ) . How-
ever, siccus is slightly smaller and much
darker than oplionoides, being black with-
out or with only faint metallic tinge while
ophonoides is always plainly greenish black.
In habits, this species resembles the pre-
ceding one (Jiamifcr mah'lieri) and often
occurs with it, under co\er on compara-
tively dry ground in more or less open
places.
Tribe OODINI
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae 5,
p. 1000 (see for synonyiiiN- and additional reter-
enees).
Ooclitac Auet. including Jeannel 1949, Coleop.
Carabicjues de la Region Malgaelu*. Part 3, p.
828.
The members ol this tribe are o\al,
Amara- or even dytiscid-like, and are black
or metallic, usually unmarked. Tlu'ir generic
classification is unsatisfactory (sec^ below.
and see Notes under Anatrichis and Oodes
hu'vissimus). However, the 13 species of
the tribe known from New Cuin(>a obvi-
oush' inc-huU' no striking ciKk'niic genera
and, although diNcrse, (lu'\ are less so than
the oodinc\s ol tlu' Oiiental lu'gion or Aus-
tralia. The Oriental SysfoU)cranius, Ilolco-
coleus, Sinu)us (see under Oodes hievis-
sinuis), etc., and th(> Australian Co])tocarpus
do not reach New (hiinea.
The Carabid Beetles of New Guinea • Darlington 31
The presence or absence of a basal elytral
margin is a useful key character in this
tribe but must be determined with care.
The basal margin is a fine, sharph' marked
line, impressed or formed by a slight eleva-
tion of the elytral surface in front of the
margin, and usually ending inwardly op-
posite the bases of the 3rd striae or 3rd
intervals. It is distinct from the basal de-
pression of the elytra that fits under the
base of the pronotum. It is best examined
under diffused light, for a sharply focused
spotlight makes reflections that simulate a
margin where none is present.
Presence or absence of a small seta-bear-
ing puncture on the posterior edge of the
pronotum on each side near the basal angle
is another useful key character that must
be determined \\'ith care. The setae are
sometimes very small and weak and easily
broken off. The punctures may then be
hard to detect even in clean specimens and
undetectable in dirty ones. Sometimes these
setae and punctures vary within single
species (see Oodes siamensis).
Clean specimens of Anotrichis pusilla,
Oodes exiguus, and O. piceus can be seen
to have a small anterior puncture over each
eye, with or without a small, weak seta.
These punctures are lacking in all other
New Guinean Oodini. This suggests an
actual relationship among the 3 species
named, which is supported by the arrange-
ment of labral setae and by a similarity
of body form, and this in turn suggests
that the conventional distinction between
Anatrichis and Oodes is unnatural. How-
ever, I cannot recharacterize these 2 widely
distributed genera on the basis of the few
I species that occur in New Guinea.
Most oodines are ac[uatic or subaquatic,
living in vegetation or among dead leaves
in or close to standing water, but O.
laevlssimus Chaudoir and probably also the
I 2 related species described below (i.e., the
laevissimus group) live in leaf litter on the
floor of rain forest. This is the habitat of
some Copiocarpus in Australia and of cer-
tain other oodines elsewhere. State of wings
is correlated with habitat. The wings are
fully developed in all known New Guinean
Oodini except the Oodes laevissimus group,
in which the wings are apparently atrophy-
ing as the group leaves subaquatic habitats
and invades terrestrial leaf litter. At the
same time the group is apparently begin-
ning to evolve local flightless species in
different places in New Guinea.
Key to Genera of Oodini of New Guinea
1. Size .small (c. 5 mm); clypeus without seta-
lieaiing punctures; $ front tarsi only slightly
dilated (p. 31) _.__ Anatriclii.s
- Size usually larger; // size small, clypeus
with seta-bearing puncture on each side and
c^ front tarsi wider ( p. .32 ) Oodes
Genus ANATRICHIS Leconte
Leconte 1853, Trans. American Philosophical Soc.
10, p. 391.
Chaudoir 1882, Ann. Soc. Ent. France (6) 2, p.
318.
Sloane 1910, Proc. Linnean Soc. New South Wales
35, pp. 442, 443 (the Australian species).
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae 5,
p. 1003 ( see for synonymy and additional refer-
ences ) .
Diagnosis. Very small Oodini; clypeus
and posterior margin of pronotum without
seta-bearing punctures; labrum with clump
of 4 setae at middle and 1 separate seta each
side; 6 front tarsi typically only slightly
dilated, but variable (see Notes below).
Description. None required here.
Type species. Oodes minutus Dejean, of
North America.
Generic distribution. India and Burma
to Australia; tropical and warm temperate
America.
Notes. The characters and limits of this
genus are doubtful, as suggested in discus-
sion under the tribe (above).
Authorities disagree about the c^ front
tarsi of Anatrichis. Leconte says 4 segments
are slightly dilated and spongiose (with
dense squamae) below. Chaudoir says 3
segments are thus modified. And Sloane
says only 2 segments have squamae. In fact,
different species differ in this respect, and
minor variations of S tarsi may even occur
32 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
within single species (see Notes under A.
pimUa, below ) .
Anafrichis pus'illa Sloane
Sloane 1910, Pioc. Linnean Soc. New South Wales
35, p. 443.
Description ( diagnostic characters only ) .
Small, narrow; pronotum usually with an
almost punctiform impression each side
near base, but these impressions variable
and sometimes almost absent; elytra 7-
striate, striae punctulate; i tarsi slightly di-
lated, with 2 or 3 segments squamulose be-
low ( see following Notes ) ; other characters
given in preceding Key to Genera; length c.
5 mm.
Types. Described from 2 specimens
taken by Sloane near Kuranda, North
Queensland, Australia, June 1906. I here
designate as lectotype the single specimen
now in the Sloane Collection at Canberra.
It is labeled "Kuranda, Q., T.G.S., 6.06" and
"Anatrichis pusilla SI., Id. by T. G. Sloane '
( seen ) .
Occurrence in New Guinea. Papua: 5,
Dobodura, Mar.-July 1944 ( Darlington ) ; 9,
Oro Bay, Dec. 1943-Jan. 1944 (Darling-
ton); 3, Lake Daviumbu, Flv R-, Sept.
11-20 and 21-30, 1936 (Archbold Exp..
A.M.N.H.); 1, Modewa, Modewa Bay, 0-50
m, "No. 17," Dec. 14, 1956 (L. J. Brass,
Fifth Archbold Exp., A.M.N.H.). N-E.
N. G.: 1, Aitape, Aug. 1944 (Darlington).
West N. G.: 24, Ilollandia, July-Sept. 1944
(Darlington); 2, Sarmi, W. of Ilollandia,
July 20-23, 1959 (T. C. Maa, Bishop Mus.);
3, Maffin Bay, Aug. 1944 (Darlington).
Notes. I have a scries of ))UsilJa from
North Queensland, Austraha: from Cairns
(near the type locality), south to Cardwell,
and north to Silver Plains halfway up the
C^ape York peninsula. Specimens from New
(iuinea match Australian ()n(\s well.
A. ptisilla is similar to and may represent
Anatrichis indica (^haudoir of India, and I
ha\'e a related species Iroin Le\ te in the
Philil)pines.
I'hr narrowly dilated ,^ front tarsi seem
to have either 2 or 3 segments squamulose.
I cannot determine whether this is primarily
individual variation, or whether the squamae
are worn off the 3rd segments in some
individuals, or whether the squamae are
sometimes pressed against the soles of the
3rd segments and therefore almost un-
detectable.
Genus OODESBonelli
Bonelli 1810, Observations Ent. 1, table synoptique,
Mem. Acad. Sci. Turin 18, pp. 21-78.
Sloane 1910, Proc. Linnean Soc. New Soutli Wales
35, p. 442 (Australian species).
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae 5,
p. 1006 (see for synonymy and additional refer-
ences ) .
Andrewes 1940, Proc. R. Ent. Soc. London (B)
9, pp. 203 ff. (key to species of India, Burma,
etc. ) .
Jeannel 1942, Faune de France, Coleop. Carabitiucs,
Part 2, p. 980.
Diaii,nosis. No satisfactory one available.
For practical purposes all Ne\^' Guinean
members of the tribe except Anatrichis
pusilla are assigned to Oodes.
Description. None required here.
Tyjx' species. Carahus liclopioidcs Fab-
ricius, of Europe.
Generic distribution. Most of the warmer
parts of the world, but few or none in
South America.
Notes. For comments on classification,
habitats, and state of \\ings see tribe Oodini.
above.
Kiiv TO Species of Oonr.s of New Cuixea
\. Labruni with compact clump of 4 (or
fewer) .setae at nn'ddle and 1 separate seta
each side 2
- Lal)rnni with 6 or 4 separate setae (if 4,
2 adcbtional uu'nutc setae usually present
close to.uethi'r near middle) 3
2. EKtra with 7th striae almost oliliterateil;
length c. 8 mm (p. 33) ))iccu\
- EKtra with 7th striae well impressed;
length r. .") mm {p. 33) cxii^tdis
3. CKpeus without seta-bcariiisi; punctures 4
- Chi^eus with seta-bearing pmnlure each
side - 7
4. Prothorax without basal setae or punctmcs
(p. 33) - nil
Prolliorax witJi seta or small puncture on or
near posterior edue eat'h side ( hicvi.ssitiius
group) 5
The Carabid Beetles of New Guinea
Darlington
33
5. Elytra with hasal margin sharply defined
( see under tribe Oodini ) ; inner wings
full or nearly so (p. 34) laevissimti.s
- Elytra with basal margin obsolete; inner
wings vestigial 6
6. EUtral striae distinctly impressed; pos-
terior pronotal punctures on dorsal surface
at base (West N. G. ) (p. 34) rossi
- Elytral striae reduced to fine superficial
lines; posterior pronotal punctures on basal
edge of pronotum ( N-E. N. G. ) ( p. 35 ) _„_
wiboni
7. Clypeus margined anteriorly, with setae
almost in the angles (p. 36)
siamcnsis vulsus
- Clypeus not margined, with setae behind
the angles 8
8. Prothorax with seta or small puncture each
side on posterior edge 9
- Prothorax without such setae or punctures;
(form stout, convex; c$ luiddle tibiae bent
near base; length c. 13.5-15.0 mm) (p.
36 ) denisonensis
9. Elytra with basal margin sharply defined
(see under tribe Oodini) (p. 36) siccus
- Elytra with basal margin obsolete 10
10. Metepisterna sparsely, vaguely, or not
punctate (p. 37) par
- Metepisterna closely punctate 11
11. Form average, prothoracic width/length c.
1.65 (p. 37) crihristernis
- Form slender, prothoracic width/length 1.44-
1.49 (p. 38) longior
Oodes piceus Nietner
Nietner 1856, J. Asiatic Soc. Bengal 25, p. 526.
Andrewes 19.30, Cat. Indian Insects, Part 18,
Carabidae, p. 238.
1940, Proc. R. Ent. Soc. London (B) 9,
p. 205 (in key).
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae 5,
p. 1010.
Description (for recognition only). A
narrowly oval, convex Oodes with elytra 6-
striate (7th striae absent or faint) and
other technical characters given by An-
drewes (1940); length c. 8 mm. See also
preceding Key and following Notes.
Type. From Colombo, Ceylon; in Stettin
Mus. (not seen).
Occurrence in New Guinea. Papua: 1,
Oro Bav, Dec. 1943-Jan. 1944 (Darlington).
West N. G.: 1, Hollandia, 250 ft., May
1945 (H. Hoogstraal, Chicago Mus.), in
rain forest.
Notes. At the British Museum in 1947-
1948, I compared Oodes piceus with wester-
manni Laferte as identified by Andrewes,
and could find no significant external dif-
ferences except in form of S front tarsi,
which are wider in westermanni. The New
Cuinean specimens are 9 9 , so their as-
signment to piceus is tentative. Oodes
piceus has been recorded from SE. Asia,
Sumatra, Java, the Philippines, and Cele-
bes. O. westermanni occurs in the same
general area.
Oodes exiguus Andrewes
Andrewes 1933, Ent. Monthly Mag. 69, p. 56.
pijginaeus Andrewes 1936, Treubia 15, p. 218
(name used in error for exiguus).
Description ( for recognition only ) . Very
small, size of Anatrichis pusilla but differing
as noted below; see also preceding Key to
Species; length c. 5 mm.
Types. A S from Sumatra, in Deutsches
Entomologisches Mus. (not seen); a 9
"cotype" in Andrewes Coll., British Mus.
( seen ) .
Occurrence in New Guinea. Papua: 71,
Dobodura, Mar.-July 1944 (Darhngton);
12, Oro Bay, Dec. 1943-Jan. 1944 (Dar-
lington). West N. G.: 23, Hollandia, July-
Sept. 1944 (Darlington); 6, Maffin Bay,
Aug. 1944 (Darlington).
Notes. The known range of this species
is now Sumatra (the types), Leyte Is. in
the Philippines (Darlington, M.C.Z.),
Morotai Is. in the Moluccas (Darlington,
M.C.Z.), and New Guinea. It is not known
in Australia.
This small oodine differs from Anatrichis
pusiUa as follows: form wider; only 1 seta
over each eve (2 in pusilla); mandibles
longer, straighter; clypeus with seta-bearing
punctures; elytra with striae not punctulate;
c? front tarsi wider (2nd segments c. long
as wide), with 3 segments squamulose
below. Both A. pusiUa and O. cxii^uus have
elytra with humeri dentate and 3rd inter\'als
2-punctate.
Oodes nil n. sp.
Description. Form (Fig. 11) average, mod-
erately convex; black, appendages slightly
34 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
riifescent; moderately shining, whole upper
surface with microsculpture of fine c. iso-
diametric meshes and also fine punctulation.
Head 0.50 and 0.51 width prothorax; labrum
6-setose, the 2 middle setae small and close
together; clypeus not margined, without
setae; only 1 (posterior) seta over each
eye; front irregular but scarcely impressed;
mentum tooth triangular, not distinctly
emarginate. Prothorax: width length 1.67
and 1.62; base aj^ex 1.75 and 1.80; disc with
fine middle line but transverse and baso-
lateral impressions slight and poorly de-
fined; posterior edge without setae. Elytra:
width elytra /prothorax 1.05 and 1.06; basal
margin present; humeri not dentate; striae
lightly impressed, finely punctate; intervals
nearly flat, 8th wide at base, 3rd 2-punctate.
Inner wings full. Lower surjaee: prosternal
process weakly or not margined between
coxae; sides of body including metepisterna
extensively and closely punctate. Secondary
sexual charaeters: 9 with 1 seta each side
last ventral segment; 6 unknown. Measure-
ments (types); length 10.5-11; width 3.3-
3.4 mm.
Types. Holotype 9 (M.C.Z., Type No.
31,555) from Dobodura, Papua, Mar.-July
1944 (Darlington); 1 9 paratype from Oro
Bay (near Dobodura), Dec. 1943-Jan. 1944
(Darlington).
Additional material. One 9 , Maffin Bav,
West N. G., June 1944 (E. S. Ross, CaH-
fornia Acad.).
Measured specimens. The holotype and
paratyp(\
Notes. For distinguishing characters of
this species see preceding Key to S))ecies,
and Notes under Codes siccus (p. 37).
The specimen from Maffin Bay is larger
than the types (length r. 12.5 mm) but has
the same technical characters.
Oodes laevissimus Chaudoir
Cliauiloir 1882, Aim. Soc. Eiit. France (6) 2, p.
361.
Andrpwes 1924, Ann. Mag. Nat. Hist. (9) II,
p. 588 ( Simons) .
Description f for recognition onl\). I'orm
parallel, depressed; strongly shining; elytra
lightly striate; for technical characters see
preceding Key to Species; length c. 11.5-
12.5 mm.
Types. From Fly R., presumably Papiia,
collected by D'Albertis; the actual type {t.
Andrewes) in Oberthiir Coll., Paris Mus.
(not seen).
Occurrence in New Guinea. Papua: Fly
R. (the types); 22, Dobodura, Mar.-Julv
1944 (Darlington); 1, Kokoda, 1200 ft. (366
m), Aug. 1933 (Cheesman). N-E. N. G.:
19, Aitape, Aug. 1944 (Darlington); 7,
lower Busu R., Huon Pen., Apr. 4, Mav 13,
1955 (E. O. Wilson, M.C.Z.); 2, Erima,
Astrolabe Bay, 1897 (Biro); 1, Sattelberg
(British Mus.); 2, Wareo, Finschhafen (Rev.
L. Wagner, S. Australian Mus.).
Notes. This distinct species is probabl\-
confined to New Guinea, perhaps to the
eastern and central part of the island. An-
drewes referred it to the genus or subgenus
Simous, but I think this was a mistake.
Simous seems to be a natural group of about
9 known species confined to the Oriental
Region including Sumatra, Java, and Borneo,
and characterized by a very short, broad
labrum and a broad, emarginate mentum
tooth. Oodes laevissimus has the labrum
narrower, the mentum tooth narrowcM- and
scarcely emarginate.
The wings in some individuals of this
species look fully developed and fit for
flight but in others they appear slightly
reduced (but still nearly full) and unfit
for flight. It is doubtful if an\- indixiduals
realK' lly. I ha\-e seen none Irom light
traps.
Although all other New Ouinean Oodini
that I have collectt^d are ac^uatie or semi-
acjuatic-, this one is not associated w ith open
water but lives among dead leaves on the
lloor of rain forest. This is probabl\- also
the habitat ol the two related ionns de-
scribed below .
Oodes rossi n. sp.
D('scri))lioii. I'^oiin as in Figure 12. c. as
Idci'issimus. subparallcl. rathc-i" depress(>d;
The Carabid Beetles of New Guinea
Darlington
35
black; tarsi, antennae, and mouthparts more
brownish; moderately shining, entire upper
surface with very fine c. isodiametric micro-
sculpture but without or with only in-
distinct punctulation. Head 0.52 width
prothorax; labrum 6-setose; clypeus not
margined, without setae; 1 (posterior) seta
over each eye; frontal impressions distinct
but poorly defined; mentum with mod-
erately broad c. truncate tooth. Prothorax:
width ^length 1.66; base /apex 1.69; disc
flattened especially posteriorly, with middle
line (and superficial irregularities) but no
other distinct impressions; 1 well impressed
seta-bearing puncture on each side on dorsal
surface just before basal edge. Elytra:
width elytra prothorax 1.09; basal margin
obsolete; humeri not dentate; striae slightly
impressed, faintly punctulatc; intervals
nearly flat, 8th wide at base, 3rd with
2 inconspicuous dorsal punctures. Inner
wings atrophied, reduced to narrow strips c.
Vs long as elytra. Lower surface: prosternal
process not margined between coxae;
metepisterna (and rest of lower surface)
virtually impunctate. Secondary sexual char-
acters: S front tarsi moderately dilated (2nd
segments slightly wider than long), 3 seg-
ments densely squamulose below; 6 with 1
seta each side last ventral segment; 9 un-
known. Measurements: length 14; width
5.9 mm.
Type. Holotype i ( California Acad. )
from Maffin Bay, West N. G., June 14,
1944 (E. S. Ross); the type is unique.
Notes. This species has probably dif-
ferentiated locally, from /f/ei;/.s.s/r7H/.S'-like
stock, by atrophy of the wings, obliteration
of the basal elytral margin, and other small
changes.
Oodes wilsoni n. sp.
Description. Form (Fig. 13) and char-
acters of the preceding species ( rossi ) ex-
cept as follows. Head 0.51 width prothorax.
Prothorax: width/length 1.72; base/apex
1.78; basal seta-bearing punctures on (not
before) basal edge of pronotum. Elytra:
width elytra prothorax 1.06; striae very
fine, superficial. Inner wings reduced to
vestiges c. V-i long as elytra. Secondary
sexual characters: S unknown; 9 with 1
seta each side last ventral segment. Mea-
surements: length 14; width 5.9 mm.
Type. Holotype 9 (M.C.Z., Type No.
31,556) from Ebabaang, Mongi watershed,
Huon Pen., N-E. N. G., 1300-1400 m, Apr.
16-18, 1955 (E. O. Wilson); the type is
unique.
Notes. This is apparently another local-
ized flightless species derived from laevis-
simiis-like stock.
(Oodes siamensis Chaudoir)
Chaudoir 1882, Ann. Soc. Ent. France (6) 2, p.
358.
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae 5,
p. 1011.
/.s.s!/.s' Andievves 1931, J. Federated Malay Mus. 16,
pp. 434, 444, fig. 4 (new synonymy).
alesi Jedlicka 1936, Acta Soc. Ent. Czechoslovakia
33, p. 66 (new synonymy).
Description ( for recognition only ) . Form
average, depressed; black; clypeus mar-
gined, with setae in angles; see siamensis
vulsus in preceding Key to Species of Oodes,
but note basal seta-bearing punctures of pro-
notum usually present in typical siamensis
( see Notes below ) ; length c. 8-9 mm.
Types. Of siamensis, from Bangkok,
Thailand; in Oberthiar Coll., Paris Mus.
( not seen ) . Of issus, from Brunei, Borneo;
in Andrewes Coll., British Mus. (seen). Of
alesi, from Mt. Makiling, Luzon; in British
Mus. ( seen ) .
Occurrence in New Guinea. Represented
only by the following subspecies.
Notes. The synonymy suggested above
is based on examination of the types of
issus and alesi at the British Museum, and
comparison with many specimens from
other localities. They seem to represent
one variable species which is widely dis-
tributed in SE. Asia, Sumatra, Borneo,
the Philippines, New Guinea, and New
Britain, and presumably intervening is-
lands too.
My single specimen of siamensis ( issus )
from Borneo has distinct basal pronotal
36 Bulletin Museum of Comparative Zoology, Vol. 137, Xo. 1
setae on both sides, but they rise from
scarcely detectable punctures that could
hardly be seen if the setae were missing.
Some, but perhaps not all, of my specimens
of this species (alesi) from Leyte have
these setae present too. However, I have
carefully examined both sides of all 36
specimens of the species from New Guinea
and 16 from New Britain, and can find no
trace of basal pronotal setae or punctures
in any of them. This gives a basis for
separating the New Guinea-New Britain
population as a geographical subspecies
(below). First, however, I have had to
discuss siamenms as a whole, in order to
establish the synonymy and distribution of
the species.
Oodes s'tamensis vulsus n. subsp.
Description. Similar to sianieiisis scnsu
stricto (above) but without seta-bearing
punctures on basal edge of pronotum.
Head 0.51 and 0.51 width prothorax. Pro-
thorax: width/length 1.57 and 1.63; base/
apex 1.81 and 1.84. Elytra: width elytra
prothorax 1.08 and 1.07. Measurements:
length c. 8-9; width 3.3-3.7 mm.
Types. Holotype S (M.C.Z., Type No.
31,357) and 13 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington). Ad-
ditional paratypes from West N. G.: 20,
ilollandia, July-Sept. 1944 (Darlington); 1,
Maffin Bay, Aug. 1944 (Darlington); 1,
"Neth. New Guinea," Oct. 20, 1944 (T.
Aarons, California Acad.).
Measured sj)eeiniens. The i holotype and
1 9 paratype from Dobodura.
Notes. This subspecies occurs also in
New Britain ( 16, Cape Gloucester, Dar-
lington, M.C.Z.).
Oodes denisonensis Castelnau
Castelnau 1867, Notes on .Vuslialiaii C^olcoptcra, p.
64.
Sloaiic 1910, Proc. Liimcan Soc. N(nv .South Wales
35, pp. U5, 447.
Csiki 1931, Cok'op. Cat., Carabidac, Haipalinac 5,
p. 1007.
Description (for recognition only). Form
broad, convex; for technical characlcM-s see
Sloane's (1910) key, and preceding Key to
Species of Oodes of New Guinea; length
c. 13.5-15 mm.
Type. From Port Denison (probably
near Bowen, Queensland), Australia; pres-
ent location of type unknown (not seen).
Occurrence in New Guinea. Papua: 1
9 , Rouku, Morehead R., Apr. 1962 {W. \\.
Brandt, C.S.I.R.O.). West N. G.: 1 9,
Merauke (south coast), sea level. Mar. 28,
1955 (L. D. Brongersma, Leiden Mus.).
Notes. The distinctixe characters of
denisonensis are based on the c5 . The two
9 9 from New Guinea agree well in non-
sexual details with specimens from Queens-
land, Australia (from Gayndah, Rockhamp-
ton, Townsville, and Kuranda),
Oodes siccus n. sp.
Description. Form and convexity average;
black, basal angles of prothorax and ap-
pendages slightly more reddish; moderately
shining, whole upper surface finely c. iso-
diametrically microreticulate and iiunctu-
late. Head 0.52 and 0.51 width pro-
thorax; labrum 6-setose; chpeus not mar-
gined, with 1 seta-bearing puncture each
side well behind angle; 1 (posterior) seta-
bearing puncture o\er each eye; front
weakly convex, scarcely impressed an-
teriorly; mentum with rounded-triangular
tooth. Prothorax: width length 1.59 and
1.68; base apex 1.82 and 1.83; disc with
fine middle line, vague wide baso-lateral
impressions, and seta-b(>aring jnmcture each
side on basal edge inside angle. Elytra:
width elytra j^rothorax 1.08 and 1.09; base
margined; humeri not dentat(>; striae mod-
erateK- impressed, laintK punetulate; in-
terxals slightK' convex, 8th wide at base,
3rd 2-punetale. Inner uiii'j^s lull. Lower
surface: prosternal process not margined
between coxae; sides of body iiieluding
metepistcMiia closely punctate. Seco)id(iry
sexual eharaclers: S w itli Ironl tarsi slightly
narrower than usual (2nd segments slightK
longer than wide), with usual 3 s(\gments
densely squamulose; i with 1, 9 2 setae
each side last \("ntr;il setiment. Measure-
The Carabid Beetles of New Guinea • Darlington 37
mcnts: length c. 10-11; width 4.0-4.5 mm.
Types. Holotype $ (M.C.Z., Type No.
31,358) and 12 paratypes from Dobodura,
Papua, Mar.-July 1944 ( DarHngton ) ; and
additional paratypes as follows. Papua: 6,
Lake Daviumbu, Flv R., Aug. 19-30, Sept.
1-10, 11-20, 1936 (Archbold Exp., A.M.
N.H.). West N. G.: 8, Hollandia, July-
Sept. 1944 (Darlington).
Measured speeitnens. The S holotype and
1 9 paratype from Dobodura.
Notes. This is similar to Oodes cribris-
ternis and its allies, but differs in having a
distinet basal elytral margin. It is similar
also to O. nil (described above) but clypeal
and basal pronotal setae are present ( absent
in nil), the elytral striae are less obviously
punctate, and the punctures of the 3rd
intervals are less impressed.
O. siccus occurs also on Morotai Is. in
the Moluccas (Darlington, M.C.Z.).
Oodes par n. sp.
Description. Form ( Fig. 14 ) more quad-
rate than usual, depressed; black, append-
ages in part more rufous; dorsal microsculp-
ture of fine c. isodiametric meshes with very
little additional punctulation. Head 0.57
and 0.59 width prothorax; labrum 6-setose;
clypeus not margined, with seta-bearing
punctiu-e each side behind angle; 1 (pos-
terior) seta over each eye; mentum tooth
entire, bluntly triangular. Prothorax: width/
length 1.57 and 1.61; base/apex 1.62 and
1.51; disc depressed, with moderate middle
line, vague transverse impressions, distinct
but poorly defined rounded baso-lateral
impressions ( sublinear in some lights ) , and
strong seta on basal edge each side inside
angle. Elytra subquadrate; width elytra/
prothorax 1.11 and 1.13; basal margin obso-
lete; humeri not dentate; striae impressed,
punctulate; intervals slightly convex, 8th
wide to base, 3rd 2-punctate. Inner icings
full. Lower sui^ace: prosternal process not
distinctly margined between coxae; sides of
body including metepisterna vaguely or not
punctate. Secondary sexual characters: S
front tarsi moderately dilated (2nd seg-
ments barely wider than long ) , 3 segments
densely squamulose; S with 1, 9 2 seta-
bearing punctures each side last ventral seg-
ment. MeaMirements: length c. 11-12;
width c. 4.6 mm.
Types. Holotype c5 (M.C.Z., Type No.
31,359 ) from Aitape, N-E. N. G., Aug. 1944
(Darlington); and 1 9 paratype, Hollandia,
West N. G., July-Sept. 1944 (Darlington).
Notes. The technical characters, espe-
cially the positions of setae and loss of the
basal elytral margin, suggest that this new
species may be allied to O. crihristernis
and longior, but j)ar differs from both in
being more quadrate and in having the
lower surface including the metepisterna
relatively smooth.
Oodes cribrisfernis Bates
Bates 1892, Ann. Mus. Civ. Genoa 32, p. 323.
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae 5,
p. 1007.
Andrewes 1940, Proc. R. Ent. Soc. London ( B )
9, p. 204 (in key).
Description ( for recognition only ) . Form
moderately slender, depressed; distinguish-
ing characters indicated in preceding Key
to Species of Oodes of New Guinea and in
Andrewes' (1940) key. Prothorax: width/
length 1.67 and 1.63; base/apex 1.63 and
1.58. Elytra: width elytra /prothorax 1.10
and 1.13. Measurements (of New Guinean
specimens ) : length 10.5-13.5; width 4.2-
5.5 mm.
Type. From Burma, in Genoa Mus. (not
seen ) .
Occurrence in New Guinea. Papua: 27,
Mihie Bay, Dec. 1943 (Darlington); 14,
Dobodura, Mar.-July 1944 (Darlington);
1, Mt. Lamington, 1300-1500 ft. (c. 400-
450 m) (C. T. McNamara, S. Australian
Mus.). N-E. N. G.: 1, Lae, Oct. 1944
(Darlington); 2, Aitape, Aug. 1944 (Dar-
lington); 4, Finschhafen, Huon Pen., 10 m,
Apr. 9-16, 1963 (Sedlacek); 1, Wau, Mt.
Missim, Morobe Dist., 880-1050 m, Feb.
8-9, 1963 (Sedlacek). West N. G.: 1,
Maffin Bay, Aug. 1944 (Darlington); 7,
Sansapor, Aug. 1944 (Darlington); 1, "Neth.
38 Bulletin Museum of Comparative Zoologij, Vol. 137, No. 1
New Guinea" without further locality, Oct.
20, 1944 (T. Aarons, California Acad.)-
Measured specimens. A pair (69) from
Dobodura.
Notes. The specimens from New Guinea
here recorded as crihristcrnis possess all
significant characters given in Bates' brief
description and Andrewes' key ( 1940 ) , but
direct comparison will be necessary to con-
firm the identification. The species (if it
is one species ) is now known from Burma,
Sumatra, and New Guinea. O. ohloniius
Castelnau of Australia seems to be allied
but is larger, duller, with finer elytral striae.
Ooc/es longior n. sp.
Description. Form as in Figure 15,
slender, depressed; black, posterior angles
of prothorax and parts of appendages (es-
pecially tarsi, palpi, antennae) ± reddish;
moderately shining, entire upper surface
with fine c. isodiametric microsculpture and
very fine inconspicuous punctulation. Head
0.56, 0.5(S, and 0.59 width prothorax; labrum
6-setose; clypeus not margined, with a
seta each side behind angle; 1 (posterior)
seta-bearing puncture over each eye; men-
tum tooth moderate, impressed or slightly
emarginate. Prothorax: width/length 1.46,
1.44, and 1.49; base/apex 1.70, 1.66, and
1.69; disc with light middle line, no distinct
sulibasal impressions but faintly impressed
each side at extreme base; 1 seta-bearing
puncture on l)asal edge each side near
narrowly rounded basal angles. Elytra:
width elytra prothorax 1.07, 1.09, and 1.09;
basal margin obsolete; humeri not dentate;
apices subangulate near suture (opposite
1st intervals); striae lightly impressed,
faintly punctulat(>; intervals slightly convex,
(Sth slightly narrower than 7th al l)ase, 3rd
2-punctate. huier ivings full. Loner sur-
face: proslernal process not distinctly mar-
gined between coxae; sides ol body below
including metepisterna rather finely, closely
punctate. Secondary .sexual characters: $
fronl larsi moderately dilated (2nd segment
c. wide as long), 3 segments densely
squamnlose b(4ow; ', with 1, 9 2 seta-bear-
ing punctures each side last ventral seg-
ment. Measurements: length c. 14-15;
width 5.2-5.4 mm.
Types. Holotype c^ (M.C.Z., Type No.
31,360) and 1 9 paratvpe from Hollandia,
West N. G., July-Sept! 1944 ( Darlington ) ;
and 1 i para type from Ambunti, Sepik R..
N-E. N. G., May 16, 1929 (Crane-Field
Mus. Pacific Exp., Chicago Mus.).
Measured specimens. The i holotype, 9
paratype from Hollandia, and S paratype
from Ambunti, in this order.
Notes. This new species has the technical
characters (setae, etc.) of crihristcrnis
( above ) but is larger and more slender ( cf.
proportions of criljri.^ternis), with elytra
subangulate at apex.
Although cri1)risfcrnis, like most Oodini,
lives in very wet places, longior may be even
more aquatic. My 2 specimens were taken
in comparatively deep water in floating
debris and vegetation.
Tribe HARPAUNI
Sloane 1898, Proc. Linnean Soc. New South Wales
23, pp. 455, 456 (key to Australian genera).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
pp. 1023-1268.
Van Emden 1953, Ann. Mag. Nat. Hist. (12) 6,
pp. 513 ff. (discussion in text).
Iliirpalkhic Jeannel 1942, Faune de France, Coleop.
Caraliic}ues, l^ut 2, pp. 57.5-584.
Harpalinae Basilevvsky 1951-1952, Ann. Mus.
Congo Beige (8), Zool., Vols. 6 and 9 (revision
of .Xfrican and Madagascan forms).
The tribe liarpalini contains a large
proportion of the common, medium-sized
(Carabidae that live on the ground in all
climates in all parts of the world. The\' are
very nmnerous in open country, fewer in
rain forest. Those that do li\c' in rain lorest
include many Lccanomcrus in eastern Aus-
tralia and most Trichotichnus in Ne\\-
Ciuinea. The tribe also contains many
smaller species that \i\v in wet places or
b(\side quiet (usually not rapidh' running)
Wider.
(Classification of genera within the Ilar-
jxilini is exceptionalh' difficult, perhaps (I
suspect) because the tribe is relatixcly re-
The Carabid Beetles of New Guinea • Darlington
39
cent in evolution and dispersal. Generic
classifications proposed for members of the
tribe in any one region fail in other regions,
and no usable classification exists for the
genera of the Oriental Region and Malay
Archipelago. The arrangement of genera
in the Junk Catalogue (Csiki, 1932) is said
to follow Schauberger, but he died without
explaining it. The classification that I am
using for the New Guinean forms ( see Key,
below) is based partly on well known
characters that are probably phylogenetic,
but nevertheless the key is partly superficial
and is intended primarily as an aid in
identification, not as a contribution to
harpaline classification.
Characters drawn from the soles of the
male tarsi are fundamental in harpaline
taxonomy but are, of course, useless in the
case of unassociated females. (The single
possible Trichotichmis that I have from
Australia is a female and therefore not
identifiable! ) And characters of the mouth-
parts, including the setae of the penultimate
segments of the labial palpi, are funda-
mental too, but are difficult to see and
understand. Even experienced carabid
specialists make mistakes in placing har-
paline genera. Bates' original placing of
Lamprophonus- and Andrewes' of Carbanus
are examples. Both genera were originally
wrongly characterized and put in the wrong
subtribes. Many of the harpalines that I
have for study from New Guinea were taken
in light traps, and this increases the dif-
ficulty. Light-trap specimens often have
moth scales adhering to and concealing
their mouthparts and tarsal soles, and scales
stuck to the tarsal soles may even counter-
feit sexual squamae.
The distribution of Harpalini over the
world has been misunderstood until re-
cently because of lack of adequate subtribal
and generic classifications, and because of
incorrect assignments of many Australian
and South American species to northern
genera, especially to Harpalus and Aniso-
dactylus. Van Emden (1953), however,
has suggested what I think are natural and
Table 1. Distribution of Principal Subtribes
OF Harpalini (after Van Emden 1953)
1. Anisodactylina: worldwide, but irregularly
distributed; genera in Australia and South America
are prolaably not directly related.
2. Harpalina, Harpali ( Harpalu.s and its im-
mediate allies ) : tliroughout Eurasia, Africa ( and
Madagascar), and North America; absent in Aus-
tralia and South America.
3. Harpalina, Selenophori: most of the world
including South America, but absent in most of
Australia (one or two Oriental genera reach just
the northern edge of Australia ) .
4. Pelmatellina: chiefly Australia and South ( and
Central) America. The genus Nemaglossa may
occur in both Australia and South America but
lias not been adequately studied.
5. Acupalpina: nearly worldwide, with some
genera very widely distributed. The members of
this subtribe are mostly small, water-loving forms
which do not compete with most members of
the other subtribes, except perhaps with small
Pelmatellina in Australia.
useful subtribes and has indicated their
distributions. His arrangement, slightly
modified, is summarized in Table 1.
This outline of harpaline distribution
(Table 1) is, of course, an oversimphfica-
tion. A more detailed study of the distribu-
tion of subtribes of Harpahni would be an
important contribution to insect zoogeogra-
phy.
Within the limits of New Guinea and
Australia, harpaline faunae overlap com-
plexly. Among larger, terrestrial Harpalini
at low altitudes, several primarily Aus-
tralian genera of subtribe Anisodactylina
(Gnathaphanus, Diciphoromerus, Hyphar-
pax) extend to New Guinea and westward
into or across the Malay Archipelago. These
genera live chiefly in relatively open
country, including open Eucalyptus wood-
land, although some of them enter rain
forest too. On the other hand, several pri-
marily Oriental genera of subtribe Har-
palina, especially Trichotichmis and other
Selenophori (but not Harpalus), reach New
Guinea and are dominant there, outnum-
bering the Australian Anisodactylina espe-
cially in rain forest. These genera either
40 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
do not extend to Anstralia or are repre-
sented there by single species on the ex-
treme northern edge of the continent (e.g.,
a CoIeoUssus on Cape York). The Austrahan
genera, chiefly in more open country, and
the Oriental ones, chiefly in rain forest, are
in part ecologically as well as geographi-
cally complementary. This pattern suggests
recent multiple invasions of the rain-
forested areas of New Guinea by Oriental
stocks and of the more open areas by Aus-
tralian stocks, but over a long period some
replacement of Australian by incoming,
competing Oriental groups may have oc-
curred.
At much higher altitudes on New Guinea
is one additional genus of Anisodactylina,
CJiydaeus, which is primarily Asiatic and
has apparently "mountain hopped" across
the Malay Archipelago. This genus does
not reach Australia.
Among smaller, water-loving Harpalini,
primarily Oriental Acupalpina are domi-
nant in New Guinea and several genera
reach the northern half of Australia, but
they decrease or disappear in southern Aus-
tralia. Their place there is taken by small
Pelmatellina {Lecanomenis), which are
ninnerous throughout Australia and a few
of which occur in New Guinea (described
in the following pages) but which are un-
known farther west in the Malay Archipel-
ago. The distributions of Oriental Acu-
palpijia and of small Australian Pelmatellina
are therefore broadly complementary too
in the Australian Region, but with wide and
complex overlapping.
Key to Geneha ok IIaiu'ai.im oi' New Guinea
1. Male front and micldle tarsi with sponge-
like soles of many densely packed, narrow
adhesive hairs 2
- Male front and nsually (not always) middle
tarsi 2-seriately scinanuilose below, or
rarely ( Lyter only) with more than 2
rows of long, narrow seales loosely ar-
ranged - 6
2. Size larger (6-lfi nun); scntellar striae
present (short in llii]>h(ir])(ix); pennltimate
segments labial palpi plnrisetose (Aniso-
dactylina ) 3
- Size smaller ( less than 5 mm in New
Guinea); scntellar striae absent; penulti-
mate segments labial palpi 2-setose
( Pelmatellina ) ( p. 45 ) Lecanameriis
3. Elytra without dorsal pimetures; wings
atrophied; (found only on high mts.) (p.
47 ) Chijdacus
- Elytra each with 1 or more dorsal pime-
tures; wings usually full 4
4. Elytra with se\eral or many conspicuous
dorsal punctures (p. 41) ,-. Gnathaphanus
- Elytra each with 1 dorsal pimcture _ . 5
5. Posterior tarsi long, basal segment much
more than 2X long as wide; hind femora
not strongly curved (p. 42) __ Diaphorovxents
- Posterior tixrsi shorter, basal segment 2X
or less long as wide; hind femora of c5
strongly curved, of 2 less so ( p. 44 )
Hyphen pax
6. Penultimate segment labial palpi with more
than 2 setae anteriorly; often larger ( 5-
11 mm) (Harpalina) 7
- Penultimate segment labial palpi 2-setose
anteriorly; often smaller (2.7-8.0 mm)
(Acupalpina) 12
7. Front tibiae broader, apex % or % wide
as tibial length (by measurement) (p.
59) Harpaloxenns
- Front tibiae narrower 8
8. Entire upper surface pubescent ( p. 48 ) __
PIdti/ntcfopus
- Upper surface not pubescent 9
9. Elytra with 3rd intervals 1-punctate or
impunctate 10
- Elytra with 3rd intervals with several
(very small) punctiues 11
10. Male front and middle tarsi with soles of
long, slender, loose (not 2-seriate) scales;
base of prosternuin and base of abdomen
not pubescent (p. 63) Lytcr
- Male front and (usually) middle tarsi 2-
seriateh' s(iuanndose; b;ise ot pronotnm
and b;ise of abdomen usually short-pubes-
cent (but see Notes under T. nwiliu.s)
(p. 48) -, - riirln>lirlmu.s
11. Last \entral segment with 2 sel;ie e;ich
side in both sexes; eh tra with sutural
angles denticulate ( in New Guinean
species) (p. 64) Coleoli.ssus
- Last ventral seguient with 1 si-ta each side
in both sexes; sutural angles not denticulate
(p. 66) Ily))luicri()u
12. Scutt'llar striae absent; anicrior iii;uginal
line of pronotnm ^\^'^'\^ and entire; length r.
7-8 nun (p. 68) .A/i<)/)/(),i,'r;i;(/.v
- Scntellar striae present; iuilerior m;irgin;il
line of pronotnm line or interruptiil ;it
middle; usualK sm;iller 13
13. Abdomen not pubescent (except for fi.xed
setae ) ( p. 69 ) Egadroina
The Carabid Beetles of New Guinea
Darlington
41
- Abdomen pubescent at least near apex 14
14. Prosternuni without long setae anteriorly
(p. 71) Stcnoloplius
- Prosternuni with several long setae an-
teriorly ( p. 72 ) Acupalpus
Genus CNATHAPHANUS Macleay
Macleay 1825, Annulosa Javanica 1, p. 20.
Chaudoir 1878, Ann. Mus. Civ. Genoa 12, pp.
476, 503.
Sloane 1900, Proc. Linnean Soc. New South Wales
24, p. 553 ( key to some Australian species ) .
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1041 (see for synonymy and additional refer-
ences ) .
Diaii.no.sis. See preceding Key to Genera
of IlarpaJini of New Guinea.
Description. None required here.
Type species. G. vidneripennis Macleay,
of Java, etc.
Generic distribution. Many species in
Australia, fewer in the Malay Archipelago
and adjacent comer of Asia, with one or
two widely distributed species reaching
India, the Philippines, and islands east to
Samoa and New Caledonia.
Notes. Some species of this genus have
very wide ranges, within the limits given
above. Of the 5 species known in New
Guinea, all are shared with Australia and
several are widespread also on the Malay
Archipelago or islands of the western
Pacific. These insects are often common in
open coimtry including grassland and open
woodland, but are not often found in rain
forest. All species of the genus that I know
are fully winged and probably fly.
Key to Species of Gnathaphanus of New Guinea
1. Elytra with intervals 3, 5, and usually 7 (at
least posteriorly) with dorsal punctures con-
spicuously impressed; (black, legs black;
elytra deeply sinuate and acuminate at apex)
(p. 41) licinoidcs
- Elytra with fewer, less impressed dorsal
punctures 2
2. Elytra with series of dorsal punctures on
outer edges of intervals 3 and ( at least pos-
teriorly) 5; (legs yellow) (p. 41) upolcnsis
- Elytra with series of dorsal punctures only
on 3rd intervals ( single punctures sometimes
present on other intervals) 3
3. Smaller ( c. 9-10 mm ) ; more shining ( es-
pecially the S ) ; piceous, legs brownish
yellow ( p. 42 ) picipes
- Larger (c. 12-13 mm); dull black or me-
tallic, legs black 4
4. Head and prothorax green, elytra cupreous
(except in discolored individuals); pos-
terior angles of prothorax distinct, bluntly
obtuse or very narrowly rounded (p. 42) ..
pulcher
- Dull black; posterior angles of prothorax
broadly rounded (p. 42) philippcnsis
Gnathaphanus licinoides Hope
Hope 1842, Ann. Mag. Nat. Hist. 9, p. 427.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1042 ( see for synonymy and additional refer-
ences ) .
Description. None required here; see
Key, above; length c. 10 mm.
Type(s). From Port Essington, northern
Australia; presumably in Hope Mus., Ox-
ford (not seen).
Occurrence in New Guinea. Papua: 24,
Dobodura, Mar.-July 1944 (Darlington); 1,
Kokoda, 1200 ft. (366 m), Sept. 1933
(Cheesman); 1, Wakaiuna, Sewa Bay, Nor-
manby Is., Jan. 1-8, 1957 (W. W. Brandt,
Bishop Mus.). N-E. N. G.: 18, Wau, 1200
m, dates in Jan., Mar., Apr., June, July,
Sept., Nov., Dec. 1961-1963 (Sedlaceks);
1, Stephansort, Astrolabe Bay, 1899 ( Biro ) ;
1, Aitape, Aug. 1944 (Darlington). West
N. G.: 1, Hollandia, May 1945 (B. Malkin,
U.S.N.M.); 1, same area, Cyclops Mts., 50-
100 m, June 22-24, 1959 (Gressitt, T. C.
Maa, Bishop Mus.), in light trap.
Notes. Tlie known range of licinoides in-
cludes northern Australia, New Britain,
the Solomons, New Hebrides, and New
Caledonia, as well as New Guinea.
Gnafhaphanus upolensis (Csiki)
Csiki 1915, Denkschriften Akad. Wiss. Wien,
Math-Nat. 91, p. 163 (Diortjche).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1044 (see for synonymy and additional refer-
ences ) .
impressipennis Castelnau 1867, Notes on Aus-
tralian Coleop., p. 100 ( in Harpalus, but not
Harpahis iiu))ressipennis Motschulsky 1844).
Chaudoir 1878, Ann. Mus. Civ. Genoa 12, p. 510.
Description. None required here; length
c. 8-9 mm.
42 BuUetin Museum of Coynparative Zoology, Vol. 137, No. 1
Types. Of impressipennis, from Rock-
hampton, Australia; in Genoa Mns. Of
upolensis, from Upolu, Samoa; in Vienna
Mils, (not seen).
Occurrence in New Guinea. Common
( 175 specimens seen ) probably throughout
New Guinea at low altitudes including
Dobodura, up to 1200 m at Wau and to
2300 m on Mt. Kaindi ( near Wau ) . Speci-
mens taken in every month.
Notes. This very^ common carabid occurs
usually in relatively open country, includ-
ing grassland and open Eucalyptus wood-
land, from the Malay Peninsula across the
Malay Archipelago to New Guinea and
Australia, east at least to the Philippines
and Samoa, and New Caledonia.
Gnafhaphanus picipes (Macleay)
Macleay 1864, Trans. Ent. Soc. New South Wales
1, p. 117 (Harpahis).
Chaudoir 1878, Ann. Mus. Civ. Gencxi 12, p. 509.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 104.3 (see for synonymy and additional refer-
ences ) .
De.<icription. None required here; length
c. 9-10 mm.
Types. From Port Denison (Bowen),
Queensland, Australia; probably in Mac-
leay Mus., Sydney (not seen).
Occurrence in New Guinea. Papua: 12,
Port Moresby, Jan., Feb., Mar., May, Aug.,
Oct., Dec. (various collectors; M.C.Z.,
British Mus., Bishop Mus., U.S.N.M., Dept.
Agr. Port Moresby), some under logs in
FAicali/))lus country, some at light; 2, Browu
K., May 22, 25, 1956 (F. J. Ford. Jr., Bishop
Mus.).'
Notes. This uortheastcrn Australian
species apparently extends only to the
southern edge of New Guinea.
Gnafhaphanus pulcher (Dejean)
Dejean 1829, Species Cleneral Coleop. 4, p. 282
( llai))altis).
Cliaiidoir 1878, .Ann. Mus. Civ. Cenoa 12, p. .505.
Csiki 1932, Coleop. Cat., Caral)idae, Harpalinae 6,
p. 1043 (.see for synonymy and additional refer-
ences ) .
Description. None required here; length
r. l.'3-16 mm.
Types. From "Nouvelle-Hollande" ( =
Australia); presumably in Oberthiir Coll.,
Paris Mus. (not seen).
Occurrence in New Guinea. Papua: 13,
Port Moresby area, various dates in Jan..
Feb., Mar., May (various collectors; Dept.
Agr. Port Moresby); 3, Bisianumu, 1600 ft.
(485 m), Feb. 12,'l966 (J. H. Barrett, Dept.
Agr. Port Moresby ) .
Notes. This Australian species apparently
reaches only the southern part of New
Guinea, perhaps only the open-wooded
Eucalyptus areas where many other Austra-
lian insects occur. It is represented on the
Lesser Sunda Islands, west to Bali, by
subspecies extrarius Schauberger.
Gnafhaphanus philippensis (Chevrolat)
Chevrolat 1841, Revue Zool., p. 221 {Amhhjgmi-
thus).
Chaudoir 1878, Ann. Mus. Civ. Genoa 12, p. 511
(as laeviceps Macleay).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1043 ( see for synon>my and additional refer-
ences ) .
Description. None required here; length
c. 12-16 mm.
Type(s). From "Manille" (Manila); in
Hope Mus., O.xford (not seen).
Occurrence in New Guinea. Papua: 1,
Kokoda, 1200 ft. (366 m). May 1933
(Cheesman); 3, Rouku, Morehead R.,
\Vest Papua, Apr. 1962 (W. W. Brandt.
C.S.I.R.O.).
Notes. Tliis species ranges from SE. Asia
to Australia, east to Philippines, but is
surprisingly scarce and perhaps localized
ill New (hiinea.
Genus DIAPHOROMERUS Chaudoir
Chaudoir 18 1.), Hull. Soc. \at. Moscow Ifi, Part
2, p. 402.
1878, Ann. Mus. Civ. Ccnoa 12. p. 17(S.
Csiki 1932, Coleop. Cat., Caral)i(lae, llarpaliuar (i,
p. 1044 (see lor additional rclerences ).
DiaL!,nosi,s. See Key to Genera of llar-
paJini of New Guinea.
Description. None rtciuircd hen-.
Type species. D. iridipennis ('haudoir, ol
.Xustralia including ("ape York (CMiaudoir
ISTS).
The Carabid Beetles of New Guinea • Darlington 43
Generic distribution. Primarily Austra-
lia, with species also on New Zealand, New
Caledonia, New Guinea, the Molueeas
(Amboina), and Timor, and with 2 New
Guinean species extending to New Britain.
Notes. Many species of this genus in
Australia inhabit open Eucah/pius wood-
land or grassland. The two species in New
Guinea occur in rain-forested parts of the
island, but I do not know their exact
habitats. The New Guinean species of
Diaphoromerus, like most Australian ones,
are winged.
Key to Species of Diaphoromerus of
New Guinea
1. Larger (8.5-10.5 mm); posterior angles of
prothorax (narrowly) rounded (p. 43) ._.
papuensis
- Smaller (6.0-7.5 mm); posterior angles of
prothorax obtusely angulate, scarcely blunted
(p. 43) papuclhis
Diaphoromerus papuensis (Macleay)
Macleay 1876, Proc. Linnean Soc. New South
Wales 1, p. 168 {Harpalu.s).
Csiki 1932, Coleop. Cat., Caral:)idae, Harpalinae 6,
p. 1043 (Gnathaphanus).
hasilewsktji Louwerens 1962, Tijdschrift voor Ent.
105, p. 139 (Gnatluiplianus) (new synonymy).
Description. None required here; length
c. 9-10 mm.
Types. Of papuensis, from Hall Sound,
Papua; presumably in Macleay Mus.,
Sydney (not seen). Of basilewskyi, from
Amboina Is., Moluccas, 70 m altitude
(A. M. R. Wegner), at light; holotype in
Louwerens Coll. (not seen), 2 paratypes
now in M.C.Z.
Occurrence in Neiv Guinea. Common
probably throughout New Guinea: 119,
from widely scattered lowland localities,
from Port Moresby and Dobodura to
Manokwari, up to 1300 m at Wau, and (SOO
m at Araucaria Camp, West N. G. Speci-
mens collected in every month except
August.
Notes. Macleay 's statement that the
third elytral interval is punctate on inner
side before apex places this species in
Diaphoromerus rather than Gnathaphanus,
and the length (4y2 lines = 9 mm) is diag-
nostic of this species in New Guinea.
Closely related species in Australia prob-
ably include mehnarius Dejean and iridi-
pennis Chaudoir. D. papuensis occurs also
in New Britain (Cape Gloucester, Dar-
lington) and the Moluccas (types of
basilewskyi) .
Diaphoromerus papuellus n. sp.
Description. Form as in Figure 16, rather
small, convex; brownish piceous, append-
ages testaceous or brownish testaceous;
moderately shining, ? scarcely duller, both
sexes with reticulate microsculpture iso-
diametric or slightly transverse on head,
more transverse on pronotum and elytra.
Head 0.69 and 0.69 width prothorax; eyes
prominent; front weakly impressed; mentum
toothed; ligula slightly shorter than para-
glossae, latter separate at apex. Prothorax
transverse-subquadrate; width/length 1.43
and 1.44; base/apex 1.41 and 1.39; sides
rounded anteriorly, nearly straight, con-
verging, sometimes slightly sinuate before
slightly obtuse but distinct and scarcely
blunted basal angles; disc formed as usual,
basal impressions sublinear, weak, mar-
gined at base but not or indistinctly punc-
tate. Elytra: width elytra prothorax 1.20
and 1.19; sides slightly sinuate before apex;
striae impressed; intervals slightly convex,
subequal, 3rd 1 -punctate on inner side near
apex. Inner wings full. Legs: 1st segment
hind tarsi elongate. Secondary sexual char-
acters: S front and middle tarsi dilated
( 2nd and 3rd segments of front tarsi slightly
wider than long, of middle tarsi narrower ) ,
with densely pubescent soles. Measure-
ments: length 6.0-7.5; width 2.0-2.8 mm.
Types. Holotype $ (British Mus.) and
6 paratypes (some in M.C.Z. , Type No.
31,361) from Kokoda, Papua, 1200 ft. (366
m), Aug. (except one specimen May) 1933
(Cheesman); and additional paratypes as
follows. Papua: 1, Kerema, May 3-9, 1959
(C. D. Michener, Bishop Mus.); 2, Kiunga,
Fly R., July 4-8, Aug. 8-10, 1957 (W. W.
Brandt, Bishop Mus.); 1, Rouku, Morehead
44 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
R., West Papua, Apr. 1962 (W. W. Brandt,
C.S.I.R.O.); 3, Yule Is. (Hungarian Na-
tional Mus.); 39, "Papua" without further
locality ( Hungarian National Mus. ) . West
N. G.': 2, Merauke, Apr. 6, 1952 (L. D.
Brongersma, Leiden Mus.); 1, same locality,
Jan. 26-Feb. 10, 1960 (T. C. Maa, Bishop
Mus.); 1, Kepi, Res. Mappi, Oct. 15, 1957
(R. T. Simon Thomas, Louwerens Coll.);
4, Wasian, Sept. 27, 1939 (R. G. Wind,
California Acad.). Also 1 paratype, Koitaki,
1500 ft. (455 m), New Guinea (diyision
unknown), Oct.-Noy. 1928 (Pemberton,
H.S.P.A.).
Measured specimen.^. The i holotype and
1 9 paratype from Kokoda.
Notes. I haye seen a specimen of this
species from Kerayat, New Britain (E. J.
Ford, Jr., Bishop Mus.).
This is eyidently a member of the
Diuphoromerus austral is group. As com-
pared with austral is itself, the present new
species has better defined posterior pro-
thoracic angles. In this character it agrees
with D. aereus Dejean, of SW. Australia,
but papueUus lacks the obyious punctation
of the base of the pronotum of aereus. As
compared with queen.slandicus Csiki (man-
dihularis Castelnau), papueUus is larger,
with more obtuse posterior prothoracic
angles.
Genus HYPHARPAX Macleay
Macleay 1825, Annulosa Javanica 1, p. 22.
Chaudoir 1878, Ann. Mus! Civ. Genoa 12, p. 496.
Sloanc 1898, Proc. Liniican Snc. New South Wales
23, pp. 458-159.
Csiki 1932, Coleop. Cat., Carahidae, Ilarpalinae 6,
p. 1051 ( se(" lor s> ii()n\ni\' and additional refer-
ences ) .
D/V/ij/fo.s/.v. See preceding Key to Genera
of llarpali)ii of New Guiiiea.
Description. None required here.
Type .species. //. lateralis Macleay ( =
dentipes Wiedemann), of Jaya.
Generic distribution. Chiefly Australia.
extending to New Zealand, and west in the
Malay Ar« •hijH'hi'io to Java and Sumatra.
Notes. See Notes under following species.
Hypharpax dentipes (Wiedemann)
Wiedemann 1823, Zool. Magazin 2, p. 54 (Harpalus).
Chaudoir 1878, Ann. Mus. Civ. Genoa 12, p. 500.
Andrewes 1919, Trans. Ent. Soe. London for
1919, p. 158.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae
6, p. 1052 (see for synonymy and additional
references ) .
Description. None required here. This
is the only species of the genus known
from New Guinea. Length c. 7-10 mm.
Type. From Java; in Copenhagen Zool.
Mus. (not seen).
Occurrence in New Guinea. Papua:
39, Port Moresby and vie, May, Sept.,
Oct., Dec. (yarious collectors; M.C.Z., Brit-
ish Mus., Bishop Mus., Dept. Agr. Pt.
Moresby), some under logs in Eucalyptus
country, some at light; 6, Yule Is., Nov. 7
and 16, 1933 (R. V. Oldham, British Mus.);
2, same locality (Fry Coll., British Mus.);
1, Lake Daviumbu,' Fly R., Sept. 11-20,
1936 (Archbold E.xp., A.M.N.H.); 1, Rouku.
Morehead R., West Papua, Mar. 1962 (W.
W. Brandt, C.S.I.R.O.). N-E. N. G.: 3, Lae
and yic. Mar. 1963, Aug. 1964 (Sedlacek);
18, Sum-Sum, 64 km N. of Wau, 580 m,
Feb. 15, 1963 (H. W. Chssold, Bishop Mus.);
4, Bulolo, 720 m, Aug. 13, 19, 24, 27, 1956
(E. J. Ford, Jr., Bi.shop Mus.), 2 of these
taken in light trap; 1, Wau, 1200 m. May
1-15, 1962 (Sedlacek) in light trap. Also
7 specimens from Papua, "British N.
Guinea," and New Guinea without exact
localities.
Notes. The sexes of dentipes differ con-
siderably: males not only have the front
and middle tarsi dilated, with spongy soles,
but also ha\e the hind femora more or less
dentate and the hind tibiae more or l(\ss
cur\'ed. The deyelopment of the femoral
tooth and the degree of cur\ature of the
tibiae xar)' indi\idually in males from sin-
gle localities and also vary geographicalK",
and the si/e of the insect varies geograph-
icalK'. The species therefore has received
several names. The synonymy has not been
fully worked out, but my impression is
that a single \ariabl(^ spcx'ies of Hypharpax,
The Carabid Beetles of New Guinea • Darlington
45
for which dentipes is the oldest name, oc-
curs in Sumatra, Java, Celebes, and New
Guinea, and that it occurs also in NE.
Australia under the name kreffi Castelnau.
This tentative conclusion should be tested
by more rigorous study, for which I now
have neither the material nor the time.
In New Guinea this species has been
found onlv in the eastern half of the island,
especially but not exclusively in the more
open Eucalyptus country of southern Papua.
Genus LECANOMERUS Chaudoir
Chaudoir 1850, Bull. Soc. Nat. Moscow 23, Part 1,
p. 446.
Sloane 1920, Proc. Linnean Soc. New South Wales
45, pp. 132, 137 (as synonym of Nemaf!,lossa).
Csiki 1932, Coleop. Cat., Carabidae, Haipalinae 6,
p. 1058 ( as synonym of Nemaglossa ) ( see for
additional references ) .
Thenarotes Bates 1878, Cistula Ent. 2, p. 319.
Diagnosis. Small Harpalini ( length under
5 mm in New Guinea); elytra without
scutellar striae (in New Guinean species);
penultimate segments labial palpi con-
spicuously 2-setose; S front and middle
tarsi with densely pubescent soles.
Description (characters common to New
Guinean species ) . Form c. as in Figures 17,
IS; more compact and convex than in most
Acupalpina, with margins of prothorax and
elytra relatively narrow. Head: mandibles
moderately long, straight posteriorly, curved
apically; eyes not very large but almost
contiguous with sides of mouth below;
frontal impressions deep, curved, sharply
defined; mentum with triangular tooth;
ligula 2-setose, with paraglossae attached,
longer than ligula; palpi short, apical seg-
ments subconical, penultimate segments of
labial palpi 2-setose. Prothorax subquadrate
or subcordate; disc convex, median longi-
tudinal line impressed, baso-lateral impres-
sions shallow and poorly defined, surface
of disc punctate across base, almost impunc-
tate elsewhere. Elytra: humeri prominent;
basal margin entire, rounded or obtusely
subangulate at humeri; striae impressed,
entire, not distinctly punctate; scutellar
striae lacking; 3rd intervals 1-punctate on
inner edge behind middle. Inner icings
full. Lotcer surface including abdomen
virtually glabrous except for "fixed" setae.
Secondary sexual characters: i front and
middle tarsi moderately dilated, with
densely pubescent soles; 2 setae each side
apex last ventral segment in both sexes.
Type species. Of Lecanomerus, L. in-
sidiosus Chaudoir, of SW. Australia; of
Thenarotes, T. tasmanicus Bates, of Tas-
mania.
Generic distribution. Species of Lecano-
merus (sensu lato) are diverse in Australia,
less so in New Zealand, New Caledonia,
and New Guinea. For further details see
Notes, below.
Notes. The supposed identity of Lecano-
menis (including Thenarotes) of Australia
and NemagJossa of Chile is doubtful.
Sloane ( 1920 ) , who suggested it, did so
without what would now be considered
critical study, and I have not been able
to make the comparisons necessary to con-
firm it. I shall therefore tentatively treat
Lecanomerus as distinct from NemagJossa
and confined to the Australian Region. The
genus does not have an "Antarctic" distribu-
tion pattern. Species are numerous and
diverse along the whole eastern edge of
Australia north to Cape York. Five species
are reported from Tasmania (Sloane), but
4 of them occur on the Australian mainland
too, and the 1 species endemic to Tasmania
is not much differentiated.
The 3 small, compact Lecanomerus found
in New Guinea resemble, but are specifi-
cally distinct from, certain unidentified
species that I found common on the Cape
York Peninsula of Australia in 1958. The
New Guinean forms occur in rain-forest
areas, not in Eucahjptus country. They
probably live among dead leaves and under
vegetation on the ground near standing
water or perhaps sometimes in leaf litter on
the floor of rain forest. However, I did not
distinguish them in the field and cannot be
sure of their habitats.
46 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Key to Species of Lecanomervs of
New Guinea
1. Prothorax narrowly subcortlate ( width /length
1.22 and 1.18); '(Hollandia, West N. G.)
( p. 46 ) angustior
- Prothorax wider; { Papna ) 2
2. Brown: slightly narrower {cf. proportions in
Description.^); piinetation of base of pronotnm
discontinnous, with middle of base virtnalK'
impnnctate (p. 46) incdius
- Black; relatively sHghtly wider; punctation
somewhat irregular but c. continuous across
base of pronotnm (p. 46) latior
Lecanomerus angustior n. sp.
Description. With characters of genus;
fomi (Fig. 17) narrowly compact; color
brownish piceous, prothoracic and elytral
margins and suture usually slightly rules-
cent, appendages testaceous; moderately
shining, reticulate microsculpture faint,
slightly transverse on front and on pronotal
disc, more transverse on elytra. Head 0.69
and 0.69 width prothorax; eyes smaller than
average, genae slightly rounded-oblique.
Prothorax: width length 1.22 and 1.18;
base apex 1.13 and 1.15; sides weakly
rounded anteriorly, converging and usually
sinuate posteriorly before c. right posterior
angles; base and apex unmargined at least
at middle; base of pronotum punctate at
sides, scarcely so at middle. Elytra: width
elytra/prothorax 1.47 and 1.45. Measure-
ments: length 3.6-4.0; width 1.6-1.7 mm.
Types. Holotype i (M.C.Z., Type No.
31,362) and 6 paratypes all from Hollandia,
West N. G., July-Sept. 1944 (Darlington).
Meamired specimens. The .', holotype and
1 9 paratype.
Notes. See preceding Key to S))ecies for
diilerential characters. This is the western-
most known species ol the genus. It prob-
ably represents medius (below) ol eastern
New Guinea. Perhaps additional related
forms are still to be found in western West
N. G.
Lecanomerus medius n. sp.
Description. With charactc-rs of genus;
form average; color brownish i)ice()us, mar-
gins of jirothorax and elytra slighth' or not
rufescent, appendages testaceous, antennae
slightly browner except at base; shining,
reticulate microsculpture faintly indicated,
meshes scarcely distinct at 50 X. Head
0.64 and 0.65 width prothorax; eyes mod-
erate, genae short, rounded. Prothorax:
width length 1.34 and 1.34; base apex 1.26
and 1.27; sides broadly rounded except
nearly straight and converging posteriorly
to obtuse but finely denticulate posterior
angles; base and apex unmargined at least
at middle; base punctate at sides, not at
middle. Elytra: width elytra prothorax
1.36 and 1.34; humeri broadh' rounded.
Secondary .sexual characters as for genus.
Measurements: length 3.5-4.3; width 1.6-
1.8 mm.
Types. Holotype i (M.C.Z., Type No.
31.363) and 20 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington), and
9 paratypes, Oro Bay, near Dobodura, Dec.
1943-Jan. 1944 (Darlington).
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. See Notes under preceding and
following species.
Lecanomerus latior n. sp.
Description. With characters of genus;
form ( Fig. 18 ) compact, relati\ely broad;
black, appendages brownish testaceous;
shining, c. without reticulate microsculpture.
Head 0.68 and 0.67 width prothorax; eyes
slightly larger than in nwdius (above),
genae short, forming c. right angles with
neck. Prothorax: width length 1.36 and
1.39; base apex 1.24 and 1.16; sides broadl\-
rounded except c. straight and conxerging
posteriorly to obtuse but minutel\ dentic-
ulate posterior angles; apex margined but
marginal line sometimes laint at middle;
base not margined; entire base ol pionotum
punctate, but punctures sparser at middle
ol base. Elytra: width el\tra prothorax
1.44 and 1.38; humeri obtuseK' sometimes
vagueK- subangulate. Measurements: length
3.6-3.7; width 1.6-1.7 mm.
Tyj)es. Holotype 6 (M.C.Z., Type No.
31.364) and 1 9 paratope from l^obodura.
The Carabid Beetles of Ne\\' Guinea • Darlington 47
Papua, Mar -July 1944 (Darlington); and
additional paratypes from Papua as fol-
lows: 1, Bisianumu, near Sogeri, 500 ni.
Mar. 15-20, 1955 (E. O. Wilson, M.C.Z.),
taken in rain forest; 1, Kokoda, 1200 ft.
( 366 m ) , Aug. 1933 ( Cheesman ) .
Measured speeimens. The 6 holotype and
$ paratype from Dobodura.
Notes. Distinguished from the 2 pre-
ceding species by broader form, black
color, and pronotum with entire apical
marginal line and more extensive basal
punctation.
Genus CHYDAEUS Chaudoir
Chaudoir 1854, Bull. Soc. Nat. Moscow 27, Part 1,
p. 343.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1080.
Diagnosis. See Key to Genera of Harpalini
of New Guinea.
Description. None required here.
Tijpe species. C. obscurus Chaudoir, of
India.
Generic distribution. Tlie Himalayas
(Sikkim, etc.), Formosa, Sumatra, Java,
the Philippines, and New Guinea; usually
at high altitudes.
Notes. This is the clearest case I know of
an Asiatic stock of Carabidae that has
"mountain hopped" to New Guinea. All
species of the genus are generally similar
and probably closely allied. The wings of
some species have atrophied, but those of
others are still fully developed, and C.
bakeri Andrewes is dimorphically winged
at Baguio on Luzon. Flying individuals
may therefore have dispersed from moun-
taintop to mountaintop and from island
to island across the Malay Archipelago
rather recently, in terms of evolutionary
time.
Chydaeus papua n. sp.
Description. Form as in Figure 19, stout,
convex; black, legs brownish, antennae and
mouthparts irregularly brownish testaceous;
both sexes moderately shining, upper sur-
face irregularly punctulate but reticulate
microsculpture faint or absent. Head 0.76
and 0.74 width prothorax, c. as in Chydaeus
obscurus Chaudoir; mentum toothed; ligula
free at apex, truncate; paraglossae arcuate,
narrow, c. long as ligula but widely sepa-
rated from it. Prothorax broadly subcordate;
width/length 1.48 and 1.49; base apex 1.12
and 1.09; sides broadly rounded through
much of length, sinuate before well defined
c. right posterior angles; pronotum strongly
convex (more so than in obscurus), base
margined, basal impressions poorly de-
fined, surface of disc more closely and
coarsely punctate at sides and especially
base than at middle. Ehjtra: width elytra/
prothorax 1.21 and 1.25; humeri subdentate;
apices weakly sinuate; striae entire, rather
lightly impressed; 3rd intervals without
dorsal punctures. Inner wings vestigial.
Lower surface and legs without obvious
special characters. Secondary sexual char-
acters: S front tarsi moderatelv and middle
tarsi narrowly dilated, densely squamulose
below; 6 with 1, 9 2 setae each side last
ventral segment. Measurements: length
9.2-10.6; width 3.6-4.4 mm.
Types. Holotype i (M.C.Z., Type No.
31,365) and 6 paratypes from Mt. Wilhelm,
Bismarck Rge., N-E. N. G., above 10,000
ft. (above 3000 m), Oct. 1944 (Darlington),
in open country above timberline; and
additional paratypes as follows, all from
the Bismarck Rge.: 2, Mt. Wilhelm, 2800-
2900 m, July 6, 1963 (Sedlacek); 1, "No. 5,"
Piunde-Aude Camp, east slopes Mt. Wil-
helm, June 13, 1959 ( L. J. Brass, Sixth
Archbold Exp. to Papua, A.M.N.H.); 1,
Lake Aunde, 3400-3500 m, July 4, 1963
(Sedlacek); 1, Lake Sirunki, 2800-2900 m,
June 15, 1963 (Sedlacek); 6, Mt. Otto Sum-
mit, Nov. 1965 ( Dept. Agr. Port Moresby ) .
Additional material. One, Murray Pass,
Papua, 2400-2800 m, Nov. 6, 1965 (Sed-
laceks); 1 i , Camp E. of Mt. Wilhelmina,
Snow Mts., West N. G., 3600 m, Sept. 1938
( Toxopeus ) .
Measured specimens. The 6 holotype and
1 ? paratype from Mt. Wilhelm.
48 BuUctin Museum of Comparative Zoology, Vol. 137, No. 1
Notes. This geographically isolated Chy-
daeus is similar to obscurtis Chaudoir (of
Sikkini, etc.) but has a slightly wider head
and differs in other details.
The Snow Mts. specimen may represent
an independent population, distinguished
by wider prothorax and perhaps by other
characters, but more material is necessary
to decide this.
Genus PLATYMETOPUS Dejean
Dejean 1929, Species General Coleop. 4, p. 68.
Csiki 1932, Coleop. Cat., Carabidae, Ilaipalinae 6,
p. 1205 ( see for synonymy and additional refer-
ences ) .
Schanberger 1938, Arl^eiten niorphologische nnd
taxonomische Ent. 5, p. 41 (see for comments
on some species of the Malay Archipelago ) .
Basilewskv 1950, Ann. Mus. Congo Bcfge (8),
Zool.. 6. p. 141.
Dkiii,nosis. Medium-sized, dull black Ilar-
palini distinguished from all other members
of the tribe in New Guinea by dorsal surface
entirely coarsely punctate and pubescent.
Description. None required here.
Tij))c -species. P. vestitus Dejean, of
Africa.
Generic distribution. Africa, the Cape
Verde Islands, and Madagjascar; SE. Asia,
Japan, and the Malay Archipelago to the
Philippines and New (iuinea (not Aus-
tralia ) .
Notes. A single widely distributed species
of this genus reaches New Guinea.
Plafymetopus iaficeps Dejean
Dejean 1829, Species Ceneral Coleop. 4, p. 70.
Csiki 1932, Coleop. Cat., (^aral)idae, Ilarpalinae 0,
p. 1200 (see tor additional rcterenccs and for
"varieties" ) .
Description. None re([uired here. See
DiciiS.nosis of genus. Length r. 8 mm.
Type(s). From the Philippines; in
Obcrthiir Coll., Paris Mus. (not seen).
Occurrence in Netv Guinea. Gollectcnl
only in the western part of West N. (i.:
4, Biak Is., dates in Jan., Feb., Mar., Apr.
1952 (L. 1). Bronger.sma, Leiden Mus.), at
light; 2, Wong R., Feb. 9, 1957 ( 1{. T. Simon
Thomas, Louwerens Coll.), at liirht; 1,
Sorong-Doom, Feb. 9, 1957 (R. T. Simon
Thomas, Louwerens Coll.), at light.
Notes. Closely related fomis of this
genus, some treated as varieties of Platy-
metopus jlaiiJahris (Fabricius) by Csiki,
are widely distributed in SE. Asia and the
Malay Archipelago. Their taxonomy is a
problem. The problem, however, lies mainly
in the Oriental Region rather than New
Guinea, and I cannot undertake to solve it
now.
Whatever the final taxonomic arrange-
ment, it seems clear that one, dark-legged
form of Phitymetopus (all surely New
Guinean individuals are dark-legged ) has
reached New Guinea recently from the
west and may perhaps still be confined to
the western end of the island. Its absence
elsewhere in New Guinea is suggested by
the facts that members of this genus are
usually common where they occur at all
and that they fly to light, but that none
has been found in light trap material from
central and eastern New Guinea. P. hiti-
ceps has been prexiously known from Bnrn
( specimen in Andrewes Coll. ) and the
Philippines.
Besides the dark-legged indixiduals re-
corded above, I have seen two yellow-
legged ones labeled "Dor)" and "Dorey."
They were probabK' collected by Wallace
and are presumably really from ('elebes
(see Part I of the present work, pp. 330-
331). They arv probably referable to P.
sidjrugosus Schanberger (see reference cited
under genus, above ) of Celebes. This
species should not be listed from New
Guinea.
Genus TRICHOTICHNUS Morawitz
Nh)ra\vil/. 1803, Mem. Acad. Sci. St. Petersburg
(7) 0, No. 3, p. 03.
Csiki 1932, Coleop. (>at., C^araliidac, Ilarpalinae
0, pp. 1210, 1217 (see for additional refer-
ences, subgenera, and synonx'nn).
l^asilcwskx 1950, .\nii. Mus. C'ongo Beige, Zool.
0, p. 87.
Di(iii,nosis. See Key to Gei\era of IlarpaJini
of New Gui)u'(i.
The Carabid Beetles of New Guinea • Darlington
49
Description ( important characters shared
liy New Guinean species). Form of or-
dinary, medium-sized Harpalini; upper sur-
face not pubescent. Head smooth except
for deep, obhque, usually linear frontal
impressions; 1 seta over each eye; mentum
toothed; labial palpi with penultimate seg-
ments with more than 2 setae. Prothoiox
subcordate or transverse; side margins each
with 1 seta-bearing puncture, before mid-
dle; disc usually extensively punctate espe-
cially across base, with punctation finer and
usually sparser across middle. Elytra with
striae entire, impressed, impunctate; 3rd in-
tervals 1-punctate near inner edge at or
slightly behind middle ( punctures some-
times obscured or absent on one or both
elytra). Inner wings usually full, rarely
dimorphic (some populations of nigricans
and altiis). Lower sw^acc: prosternum
anteriorly with short pubescence (reduced
in mcdiiis). Legs: front tibiae with apex
less than Vi wide as tibial length; hind
tarsi moderate or long. Secondary sexual
characters: 6 front and usually middle tarsi
2-seriately squamulose; 2 setae each side
last ventral segment in both sexes. See also
Notes, below.
Type species. T. longitarsis Morawitz, of
Japan.
Generic distribution. Temperate and
tropical Eurasia and the Malay Archipel-
ago to New Guinea, etc. (probably not
Australia); eastern North America. The
genus in a broad sense, including Hyparpa-
his, occurs also in Africa (Csiki, probably
following Schauberger), but Basilewsky con-
siders llyparpalus a separate genus and
does not recognize Trichotichnus in Africa
south of the Sahara.
Notes. I have had difficulty with both
the generic and the specific classifications
of the 15 New Guinean species that I now
assign to this genus. Several of the species
might go in Lampetes {Lamprophonus) or
Carbanus, but I have not found satisfactory
characters to distinguish these genera from
Trichotichnus. However, I do not intend to
reduce them to synonymy now. They need
further study based on Oriental as well as
New Guinean forms. This study will re-
quire more time and material than I now
have.
The descriptions of species in the follow-
ing pages are brief, and allowance must be
made for individual variation, which is
surprisingly great in some characters. For
example, the punctation of the outer elytral
intervals is variable in some cases (e.g., in
mixtus). The form of the elytral apices is
sometimes variable (e.g., in denarius and
altus). And the form of the apex of the
aedeagus is surprisingly variable in some
species. I have figured it in some cases
but have usually not used it as a diagnostic
character. But see under alius and dux
(Figs. 172, 173).
All New Guinean species of Trichotichnus
are fully winged and probably capable of
flight (some of them have been taken in
light traps ), with 2 exceptions. T. nigricans,
although apparently always fully winged
at low altitudes, is dimorphically winged
on the Bismarck Range. And some pop-
ulations of T. alius include individuals with
slightly shortened and weakened wings, al-
though other individuals of this species fly.
Some Trichotichnus in other parts of the
world have atrophied or dimorphic wings.
Most or all of the common, unspotted
Trichotichnus in New Guinea probably live
on the ground in rain forest, but I did not
distinguish the different species in the field
and cannot be sure of their exact habitats.
The following Key to Species of Tri-
chotichnus of New Guinea works reason-
ably well for series of clean specimens, but
it is not perfect. I myself have had trouble
placing some single specimens. In order to
simplify identifications and reduce need for
using the key, I give the following notes for
recognizing several of the commoner, dark
( unmarked ) species.
If large (7.3-S.3 mm), rufo-piceous,
shining, and without pubescence at front of
prosternum (but some setae at apex of
50 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
prosternal process ) : see Lyter, second
genus after TrichoticJinus.
If large (8-9 mm), broad, and with
paiiiol raised 10th intervals in elytral mar-
'j^ins: denarius.
If large ( c. 8-10 mm ) , less broad, with-
out raised 10th intervals, and found at con-
siderable elevations in mountains (usually
over 1200 m ) : probably altus.
If small ( 6.5-7.5 mm ) , dark, and without
reticulate microsculpture on elytral inter-
vals: probably nigricans.
If small (6.3-7.0 mm), dark, and with
reticulate microsculpture on elytral inter-
vals: probably semimas (which lacks
squamules on 6 middle tarsi ) .
Key to Species of Trichotichnus of
New Guinea
1. Head c. % width prothorax (H/P 0.66 and
0.68); prothorax transverse with broadly
rounded sides; length 5.0-6.3 mm (p.
50 ) siraneoi
- Head c. % or more width prothorax (by
measurement); prothorax more subcordate;
usually larger 2
2. Elytra with partial raised 10th intervals in
marginal channels; (no dorsal markings;
length 8-9 mm) (p. 51) denarhi.s
- Elytra without partial raised 10th inter-
vals 3
3. Male with only anterior (not middle)
tarsi squamulose; elytral intervals micro-
reticulate; ( no dorsal markings; length 6.3-
7.0 mm) (p. .52) .semimas
- Male with middle as well as anterior tarsi
with sf|uamides; elytra often (not always)
without microrcticuhition 4
4. Elytra without subapical sutural spot or
sutural intervals pale 5
- Elytra with eonnnon subapical sutural spot
pale, or sutural intervals pale near apex 11
5. Prothorax and elytra without pale margins;
abdomen usually without (•(>)is))icu()us pale
spots or margins; femora not coii.s))icuotisly
paler than abdomen 6
- Protliorax and elytra usually with narrow
pale margins; abdomen usually with con-
spicuous pale lateral spots or margins;
femora conspicnotisUi pale 10
6. Length 6. .5-7. 5 mm; pronotum not iiiucli
depressed at sides toward base; (basal
margin of pronotum usually incomplete)
(p. 52) )u^iic(nis
- Larger, or sides of pronotum more de-
pressed toward base 7
7. Eyes slightly larger, separated from mouth
below by c. '/s diameter of an eye; apex
of aedeagus short; (length c. 8-10 mm)
(p. 53) modiLs
- Eyes slightly smaller, more distant from
mouth below; apex of aedeagus longer,
slender 8
8. Length lL.5-13.0 nun (p. ,53) dii.x
- Length 7.2-10.0 mm 9
9. Prosternal pubescence more abundant; size
usually larger (c. 8-10 mm); pronotum
more punctate, less shining (p. 54) _ (dtus
- Prosternal pui^escence usualK' scanty, but
variable; size usually smaller (7.2-8.5
mm); pronotum less punctate, more shining;
(direct comparison necessary to determine
some specimens) (p. 55) mediii.s
10. Elytral intervals not obviously microreticu-
late; abdomen with pale spots usually
largest and most conspicuous at sides of
subapical segment; (length 8.3-9.3 mm)
(p. .56) hrcnidti
- Elytral intervals microreticulate; abdomen
usually more extensively pale margined;
(length 8..5-9.4 mm) (p. .56) ohscurus
11. Length usually 7.6-8.5 mm ( rarely slightly
smaller); subapical sutural pale spot dis-
tinct, reaching 3rd intervals (p. 57) _
guttuhi
- Smaller; sutural pale spot \ariable, some-
times smaller or less distinct 12
12. Prothorax wide at base ( base, ape.\ 1.34);
(length 7.5 nun) (p. 57) mongi
- Prothorax narrower at base 13
13. Sides of elytra (intervals 8, 9) rugose-
punctate; (length 5.3-6.8 mm) (p. 58) __
semirugosiis
- Sides of elytra not rugose, although some-
times punctulate 14
14. Length 5.8-7.5 mm; pronotum depressed
at sides toward base; ( and see Notes under
this species) (p. 58) mixtu.s
- Length .5. .3-5. 8 mm; sides of pronotum
scarcely deprcssi'd (p. 59) dclicatu.s
Trichotichnus sfraneoi (Louwerens)
Louwercus 1962, Tijdschrift \c)or Ent. 105, p. 1 12,
iig. 7 {C(irhanu.s).
Dc.scri))lion. With charactt^rs of genus;
small, form (Fig. 20) broad; brownish
black, sides ol piouotum and c']\ tr;i \aguely
translucent, lower snrlacc and appendages
more rufous; elytr;i fainth- iridescent but
not distinctK' microreticulate (at 50x).
Head small, 0.66 and 0.68 width prothorax;
eyes ku-gc. s('p;uated from month below b\
The Carabid Beetles of New Guinea • Darlington
51
c. ^i-j or less width of an eye. PiotJiomx
transverse with broadly rounded sides;
width length 1.59 and 1.59; base/apex 1.41
and 1.39. Elytra: width elytra prothorax —
and 1.30. Secondary sexual characters nor-
mal for Trichotichnus. Measurements (in
New Guinea ) : length c. 5.0-6.3; width c.
2.3-2.7 mm.
Types. Holotype $ (Louwerens Coll.),
allotype, 6 paratypes all from Amboina Is-
land^ Moluccas, 70 m (A. M. R. Wegner),
at light; a paratype now in M.C.Z. (Type
No. 31,149) (holotype not seen).
Occurrence in New Guinea. Papua: 3,
Brown R., May 21, 23, 24, 1956 ( E. J.
Ford, Jr., Bishop Mus.); 1, Mts. between
Agamoia and Ailuluai, Ferguson Is., 900 m,
"No. 4," June 5-17, 1956 (L. J. Brass,
U.S.N.M.). N-E. N. G.: 13, Wau, Morobe
Dist., 1200, 1300 m, various dates (Sedla-
cek); 1, Wantoat, Finisterre Mts., 4000 ft.
(1220 m), Sept. 9, 1957 (Monroe and
Holland, Canadian Nat. Coll.); 1, Elipta-
min Vy., 1350-1665 m, June 23-30, 1959
(W. W. Brandt, Bishop Mus.). West
N. G.: 1, Hollandia, July-Sept. 1944 (Dar-
lington); 1, Cyclops Mts., Sabron, 2000 ft.
(610 m), June 1936 (Cheesman); 1, Hol-
landia area, W. Sentani, Cyclops Mts., 50-
100 m, June 22-24, 1959 (Gressitt & T. C.
Maa, Bishop Mus.), in light trap; 2, Star
Rge., Sibil, 1260 m, May 16, June 16, 1959
(Leiden Mus.), at Hght.
Measured .specimens. A pair ( i 9 ) from
Brown R.
Notes. Andrewes placed Carhanus in the
wrong subtribe of Harpalini, erroneously
considering it a member of the Acupalpina.
The arrangement of setae on the labial palpi
seems to me to place it with the Haipalina,
and I can find no positive character to
distinguish it from Trichotichnus. How-
ever, I do not intend to synonymize Car-
hanus now. It requires further study. The
name can be used for a group of small,
mutually similar species (lautus Andrewes
of Burma, flavipes Andrewes of Java,
pliilippinus Jedlicka of the Philippines, and
straneoi Louwerens of the Moluccas, New
Guinea, etc. ) that may eventually be sepa-
rated from Trichotichnus.
T. straneoi extends to New Britain and
New Ireland (specimens in Bishop Mus.).
Nothing is recorded of its habitat or habits
except that it flies to light.
Trichotichnus denarius n. sp.
Description. With characters of genus;
form slightly broader than usual; black or
piceous, appendages browner, sides of ab-
domen with small pale marks (variable);
rather shining, most of upper surface with-
out visible microreticulation (at 50x ) but
elytra silky in some lights. Head 0.77 and
0.77 width prothorax; eyes large, separated
from mouth below by e. ^o diameter of
an eye. Frothorax transverse-subcordate;
width/length 1.59 and 1.63; base/apex 1.14
and 1.10; sides converging and usually
broadly and slightly sinuate before distinct
but obtuse posterior angles; basal marginal
line fine or interrupted at middle; disc
weakly convex, moderately depressed at
sides especially posteriorly, extensively
punctate but with punctation finer and
slightly sparser at middle. Ehjtra broad;
width elytra/prothorax 1.36 and 1.36; mar-
ginal gutters wider than usual, with ir-
regular raised 10th intervals in c. middle
V3 of length; apices usually slightly dehis-
cent and c. pointed, but variable; outer
intervals (8, 9, and 10) usually slightly,
finely punctulate. Secondary sexual char-
acters as for genus. Measurements: length
c. 8.0-9.0; width 3.3-3.8 mm.
Types. Holotype S (M.C.Z., Type No.
31,366) and 111 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. Papua:
20, Kokoda, 1200 ft. (366 m), June, July,
Aug., Sept., Oct. 1933 (Cheesman); 1, same
locality, Mar. 28-29, 1956 (Gressitt); 2,
Biniguni, Gwariu R., 150 m, "No. 3," July
27-Aug. 14, 1953 (Geoffrev M. Tate,
A.M.N.H.); 5, Peria Ck., Kwagira R., 50 m,
"No. 7," Aug. 14-Sept. 6, 1953 (Geoffrey
M. Tate, A.M.N.H.); 1, Kokoda-Pitoki, 400
m, Mar. 23, 1956 (Gressitt). N-E. N. G.:
52 BiiUefin Museum of Comparative Zoology, Vol. 137, No. 1
3, lower Busu R., Huon Pen., May 4, 1955
(E. O. ^^'ilson, M.C.Z.), in lowland rain
forest; 1, Simbang, Huon Gulf, 1899 (Biro).
West N. G.: 1, Wamoro (ex Coll. G.
Hauser, British Mus.).
Measured .speeimens. The c5 holotype and
1 9 paratype from Dobodura.
Notes. The partial 10th intervals in the
elytral margins immediately distinguish
denarius. The species is very common in
eastern New Guinea ( apparently much
less so in the west), presumably in ram
forest. One of Miss Cheesmans specimens
was taken at light and so apparently were
the Peria Creek individuals, which have
scales and wing fragments of other insects
on them.
Trichofichnus semimas n. sp.
Description. With characters of genus;
form rather slender; black or piceous, ap-
pendages brown, abdomen with some ( vari-
able) small pale marks at sides; moderately
shining but elytra with transverse micro-
reticulation distinct at 50x. Head 0.79 and
0.79 width prothorax; eyes moderate, sepa-
rated from mouth below by nearly % diam-
eter of an eye. Prothorax subcordate;
width length 1.48 and 1.48; base/apex 1.18
and 1.16; sides rounded anteriorly, con-
verging and sinuate before distinct c. right
(slightly obtuse) posterior angles; basal
marginal line faint or interrupted at middle;
disc very little depressed at sides toward
base, extensively punctate except almost
impunctate at middle. Ehjlra normal;
width elytra prothorax 1.23 and 1.26; outer
intervals not distinct])' punctulate. Second-
ary sexual characters normal except only
front (not middle) tarsi of 6 scjuamulose.
Measurements: length 6.3-7.0; width 2.5-
2.8 mm.
Tijpes. Holotype i (M.C.Z., Type No.
31,367) and 19 paratypes from Dobodura,
Pajma, Mar.-July 1944 (Darlington); 4
paratypes, Kokoda, Papua, 1200 ft. (366
m), Oct., Sept. 1933 (Cheesman); 3 para-
types, same locality. Mar. 20, 28-29, 1956
(Gressitt), in light trap; 2, Xormanb) Is.,
W'akaiuna, Sewa Bav, Nov. 1-10, 1956 and
Jan. 1-8, 1957 (W. \\'. Brandt, Bishop
Mus.). N-E. N. G.: 1, Erima, Astrolabe
Bav, 1896 (Biro); 2, Madang ("Friedrich-
\^m.-hafen"), 1901 (Biro); 1, Bulolo, 730
m, Aug. 27, 1956 (E. J. Ford, Jr., Bishop
Mus.), in light trap; 1, Sum-Sum, 64 km N.
of Wan, 580 m, Feb. 15, 1963 (Sedlacek).
West N. G. : 4, Hollandia area, W. Sentani,
Cyclops Mts., 50-100, 100, 150-250 m, June
(various dates) 1959 (Gressitt and T. C.
Maa, Bishop Mus.); 2, Ifar, Cyclops Mts.,
450-500, 400-800 m, Sept. 7,' 7-9, 1962
( Sedlacek ) ; 3, Dojo, Res. Hollandia, Apr.
1957, 1958 (R. T. Simon Thomas, in Lou-
werens Coll.); 1, Maffin Bay, Aug. 1944
(E. S. Ross, California Acad.); 1, Wasian,
Vogelkop, Sept. 1939 (Wind. M.C.Z.).
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. Males of semimas are imique
among New Guinean Trichofichnus in
lacking the usual sexual squamules of the
middle tarsi. Fc^nales resemble nii^ricans
( below ) but differ by presence of reticuhite
microsculpture on the elytra. T. semimas
probably lives in rain forest, and has been
taken at light near Hollandia as wi-ll as at
Kokoda.
Trichofichnus nigricans Schauberger
SLlianhermT 1935, Eiit. An/.cimT 15, p. 34.
De.ivription. \\'ith characters of genus;
form rather small and slender; black,
appendages brownish, sides of abdomen
with or without small, often vague pale
marks; upper surlace including elytra with-
out distinct reticulate microsculpture. Head
0.76 and 0.75 width jirothorax; eyes usually
moderati- and separated Irom mouth below
by c. ':. diam(4er ol an c\c. but e\"es smaller
and more distant Irom mouth in short-
wing(>d indi\iduals hom Bismarck Range.
Prothorax subcordate. with sides slightK
or not sinuate belore usnalK well defined
but obtuse basal angles; width IcMigth 1.43
and 1.49; base apex 1. 2 1 and 148; basal
inaruinal line usualK ineomplete at middle;
The Carabid Beetles of New Guinea • Darlington
53
disc only slightly depressed at sides basally,
extensively punctate, punctation finer and
sparser at middle. Elytra nomial; width
elytra prothorax 1.27 and 1.25; outer inter-
vals not distinctly punctulate. \V/ng.s full
in lowland populations, dimorphic on Bis-
marck Range (see Notes, below). Second-
ary sexual characters normal. Measure-
ments: length c. 6.5-7.5; width c. 2.5-2.8
mm.
Types. (Holo)type (Andrewes Coll.,
British Mus.) and 2 "cotypes" all from
Sattelberg, N-E. N. G. (G. Hauser). I saw
the type in London in 1948 and made a
satisfactory comparison with it.
Occurrence in Neiv Guinea. Common
and widely distributed: 180 specimens
from localities well distributed over New
Guinea and on Woodlark and Nonnanby
Is.; most from low altitudes, but series from
Chimbu Vy., Bismarck Rge., between 5000
and 7000 ft. (c. 1500-2100 m), and from
Wau, 1200 m. Specimens taken in every
month.
Measured specimens. A pair { $ 9 ) from
Dobodura, Papua.
Notes. T. nigricans occurs also on New
Britain, and related fonns, compared by
Schauberger (1935, p. 36), are known from
Java, Sumatra, and Celebes.
This species probably occurs in rain
forest, although the Chimbu specimens
were taken in open country.
I have not examined the \\'ings of every
specimen, but all or most of those from low
altitudes are winged, and they often fly.
They have been taken in light traps at
several localities. However, of my Chimbu
series, only 3 specimens (both sexes) have
full wings, and 11 (both sexes) have the
wings reduced to thin strips c. % as long
as the elytra. I have not examined the
wings of the 41 specimens from Wau be-
cause some or all were collected in light
traps, which would select only winged in-
dividuals.
Besides the 180 specimens that I assign
to this species without much doubt, 7
specimens from various localities in New
Guinea are assigned doubtfully, because of
slight differences in various characters.
Trichotichnus modus n. sp.
Description. With characters of genus;
form ( Fig. 21 ) average, somewhat variable
( see proportions ) ; black, legs brownish, ab-
domen ± brownish (apical segment darker)
with lateral pale areas absent or not sharply
defined; shining, elytra subiridescent but
without visible reticulate microsculpture.
Head 0.72 and 0.76 width prothorax; eyes
slightly larger than usual, separated from
mouth below by c. Vs diameter of an eye. Pro-
thorax transverse-subcordate; width/length
1.43 and 1.53; base/apex 1.30 and 1.20;
sides rounded anteriorly, converging and
usually slightly sinuate before well defined
but obtuse posterior angles; basal marginal
line faint or incomplete at middle; disc
moderately depressed at sides, extensively
punctate except c. impunctate at middle.
Elytra: width elytra /prothorax 1.32 and
1.36; marginal channels narrow, without
10th intervals; outer intervals not distinctly
punctulate. Secondary sexual characters
normal; apex aedeagus short, ± hooked
dorsally. Measurements: length 8.0-10.0;
width 3.4-3.9 mm.
Types. Holotype S (M.C.Z., Type No.
31,368) and 6 paratypes all from Dobodura,
Papua, Mar.-July 1944 (Darlington).
Additional material. Papua: 1, Nor-
manby Is., Wakaiuna, Sewa Bay, Jan. 1-8,
1957 (W. W. Brandt, Bishop Mus.). West
N. G.: 1, Wissel Lakes, Tage L., 1760
m, Aug. 4, 1955 (Gressitt). These speci-
mens are assigned to modus doubtfully.
Measured specimens. The c5 holotype and
1 9 paratype.
Notes. This species resembles denarius
in size and color, but differs by lack of
partial 10th elytral intervals.
Trichotichnus dux n. sp.
Description. With characters of genus;
fonn c. average, very large; black, append-
ages dark brown, abdomen without distinct
pale areas; upper surface finely micro-
reticulate, meshes c. isodiametric on head,
54 Bulletin Museum of Comparative Zoology, Vol 137, No. 1
increasingly transverse on pronotum and
elytra. Head 0.76 and 0.77 width prothorax;
eyes separated from mouth below by c.
% width of an eye (but eyes more deeply
covered than usual ^^'ith transparent win-
dow-like material so edges not precisely
defined). Fiothorax subcordate; width/
length 1.41 and 1.46; base/apex 1.19 and
1.23; sides rounded anteriorly, converging
and straight or slightly sinuate before well
defined but obtuse posterior angles; basal
and apical marginal lines usually faint or
interrupted at middle; disc moderately de-
pressed at sides posteriorly, finely but ex-
tensively punctate or punctulate, the punc-
tation strongest basally. Elytra: width
elytra/prothorax 1.39 and 1.30; margins
narrow; outer intervals not distinctly punc-
tulate. Inner icings full (see following
Notes). Secondary .sexual characters nor-
mal; apex aedeagus as in Figure 173. Mea-
.mrements: length 11.5-13.0; width 4.4-
5.1 mm.
Types. Holotype $ (Bishop Mus.) and
4 paratvpes (2 in M.C.Z., Type No. 31,369),
from Edie Creek, 14 km S^^^ of Wau, N-E.
N. G., 1900 and 2000 m, Oct. 4-10, 1961,
and Feb. 13, 1962 (Sedlacek). Additional
paratypes from N-E. N. G.: 3, Wau, 1700,
2400 m, Jan. 9-12, Oct. 6, 1962 ( Sedlacek ) ;
6, Kepilam, 2400 and 2500 m, June 21, 20-
22, 21-23, 1963 (Sedlacek); 1, Tambul, 2200
m, May 27-June 7, 1963 (Sedlacek); 1,
Laiagam, W. Highlands, Mar. 23, 1960 (J.
H. Barrett, Dept. Agr. Port Moresby), at
light; 1, Moke, Okapa Subd(istrict), E.
Ibghlands, 6400 ft. ( 1950 m), Apr. 17, 1962
(J. II. Barrett, Dept. Agr. Port Moresby).
2, Okapa, June 12, 1964, Jan. 10, 1965
(Hornabrook).
Mea.surcd specimens. The i holotype and
1 9 paratype from Edie Creek.
Notes. The large size distinguishes this
species from all odiers of the genus in New
Guinea. All specimens of the type series
are fully winged, but most were taken in
light traps which, of course, select winged
individuals.
Trichotichnus alius n. sp.
Description. With characters of genus;
form of c. average large New Guinean
Trichotichnu.s; black or piceous, appendages
dark brownish, sides of abdomen with in-
distinct or poorly defined (variable) pale
areas; moderately shining, elytra with trans-
verse microreticulation faintly or not visible
at 50x. Head 0.76 and 0.75 width prothorax;
eyes separated from sides of mouth below
by c. % width of an eye. Prothorax sub-
cordate; width length 1.40 and 1.50; base
apex 1.15 and 1.18; sides converging and
straight or slightly sinuate before well de-
fined but obtuse posterior angles; disc mod-
erately depressed at sides posteriorly, vari-
ably but often extensively punctate, most
conspicuously so across base and least so
across middle; basal marginal line entire,
faint, or interrupted at middle (variable).
Ehjtra: width elytra prothorax 1.25 and
1.31; margins narrow, without 10th inter-
vals; apices pointed or blunted (variable);
outer intervals not distinctly punctulate.
Inner wings dimorphic on the Bismarck
Range (type series), full and strong in
some individuals, slightly shortened (but
still folded at apex) and with slightly
weakened venation in other indixiduals.
Secondary sexual characters normal; i cop-
ulatory organs as in Figure 172. Measure-
mcnts: length 8.0-10.3; width 3.3-3.9 mm.
Types. Holotype 6 (M.C.Z., Type No.
31370) and 42 paratvpes from Chimbu
Vy., Bismarck Rge., N-E. N. G., 5000-7500
ft. (c. 1500-2300 m), Oct. 1944 (Darling-
ton); and 16 paratypes, Tomba, S. .slope of
Mt. Ilagen (Bismarck Rge.), 2450 m. May
22-24, 1963 ( Sedlacek ) .
Additional material. See Notes, below.
Measured specimens. The 6 holotype and
1 9 paratype from Chimbu Vy.
Notes. T. altus and its allies {dux, above,
and medius, below) are the common moun-
tain-living Tricliotichnus of New Guinea.
These 3 species seem clearU distinct and
have differcMit, but in part o\(>rlapping,
ranges: dux, on the mountains of the
The Carabid Beetles of New Guinea • Darlington 55
Morobe area; typical alius, on the Bismarck traps, and such material usually includes
Range; and typical medius, on the Torri- only fully winged individuals and is not
celli Mts. However, I have seen many satisfactory for study of wing dimorphism,
additional specimens of alius or closely This is another reason for not attempting
related forms from other localities, as fol- a more detailed study of alius and related
lows. Papua: 9, Mt. Giluwe, 2500 m. May species with the material available now.
1 and 27, June 6, 1963 (Sedlacek); 1,
Dimifa, SE. of Mt. Giluwe, 2200 m, Oct. Trichotichnus medius n. sp.
11, 1958 (Gressitt); 1, Owen Stanley Rge.,
Goilala, Bome, 1950 m. Mar. 8-15, 1958 Description. With characters of genus;
(W. \y. Brandt, Bishop Mus.); 1, Mafulu, form average; black, appendages brownish
4000 ft. (1220 m), Jan. 1934 (Cheesman). testaceous, abdomen with or without poorly
N-E. N. G.: 270 specimens (in addition to defined pale lateral areas (variable), hind
the type series) from localities including femora not sirikino^ly paler than abdomen,
Morobe Dist.; Kratke Mts.; W. Highlands; tibiae paler than femora; shining, elytra
and (S. of the Markham-Ramu Vy.) Sala- not visibly microreticulate at 50x. Head
waket Rge; Mongi Watershed; and Huon 0.78 and 0.79 width prothorax; eyes sepa-
Pen. West N. G.: 35 specimens, from rated from mouth below by % or Vs diam-
localities including the Star Rge.; Wissel eter of an eye. Pwihorax subcordate; width/
Lakes; and Snow Mts. (Top Camp; lebele length 1.49 and 1.52; base/apex 1.18 and
Camp; Mist Camp; Baliem Camp). Most 1.14; sides converging and straight or
specimens are from altitudes of 1200 to slightly sinuate before well defined but
2700 m, but a few, from within 200 m of obtuse basal angles; fine basal marginal
sea level. Individuals have been taken in line usually complete; disc moderately de-
every month. Most specimens are in the pressed at sides basally, extensively punc-
British Mus., Bishop Mus., A.M.N.H., tate across base, punctation much finer
Leiden Mus., C.S.I.R.O. Coll., and M.C.Z. and somewhat sparser across middle and
I have restricted the type series of alius anteriorly. Elyira: width elytra prothorax
and its close relatives to specimens from 1,31 and 1.26; margins without 10th inter-
single localities or restricted areas because vals; outer intervals not distinctly punc-
the species of this group obviously vary tulate. Inner icings full. Lower sutiace:
geographically as well as individually. In flnfen'or part of prosternum with pubescence
general, specimens from north of the Mark- usually sparse (but variable). Secondary
ham-Ramu Valley average larger, those sexual characters normal. Measurements:
from south of the valley smaller, except length c. 7.5-8.5; width 3.1-3.3 mm.
that some specimens from Wau are as small Types. Holotype S ( Bishop Mus. ) and
as some of the types of medius. Two dis- 22 paratypes (some in M.C.Z., Type No.
tinct forms, a very large one (dux) and a 31,371) from Mokai Village, Torricelli Mts.,
smaller one (tentatively referred to alius), N-E. N. G., 750 m, various dates in Dec.
occur at Edie Creek, showing that the 1958 and Jan. 1959 (holotype, Jan. 1-23,
species of this group are not entirely al- 1959) (^^^ W. Brandt); and additional
lopatric. The characters, variation, and dis- paratypes, all from Torricelli Mts., as fol-
tribution of these species need more study lows: 19, Mobitei, 750 m, dates in Feb.,
than I can give them now. Mar., Apr. 1959 (W. W. Brandt, Bishop
T. alius is known to have dimorphic wings Mus. ) ; 3, Wantipi, Nov. 30-Dec. 8, 1958
(see Description) only on the Bismarck (W. W. Brandt, Bishop Mus.).
Range. The specimens in question were Additional material. Some specimens
not collected at light. Much of the other among those summarized under Trichoti-
material listed above was taken in light chnus alius may prove referable to medius,
56
Bulletin Museum of Coiupuratwe Zoologij, Vol. 137, No. 1
especially the smaller ones from south of
the Markham-Ramii Valley and from Wau.
Measured specimens. The 6 holotype and
1 9 paratype from Mokai.
Notes. This species is difficult to define
exactly. It is larger than niii,ric(U]s, with
basal marginal line of pronotum usually
entire (usually interrupted in ni'^ricans).
It is smaller than modus and (dtus, with
eyes intermediate in size. The sparseness
of pubescence on the anterior part of the
prosternum is an aid in identification, but
it is not infallible. This species is more
shining than most Trichofichnus, and ap-
proaches Lijter (p. 63) in appearance, but
the clothing of the male front and middle
tarsi of mcdius consists of 2 rows of broad
scales as usual in Trichotichnus. Neverthe-
less, a Tricliolicluius like the present one
may have been ancestral to Lytcr.
Trichotichnus brandti n. sp.
Description. With characters of genus;
form average; black or piceous, not marked
above except lateral margins of elytra and
sometimes of prothorax narrowly incon-
spicuously rufesccnt or translucent; reddish
piceous below with epipleurae and narrow
maigin of abdomen paler, the pale marks
usually widest and most conspicuous at
sides of subapical ventral segment; ap-
pendages brownish testaceous, femora con-
spicuously paler than abdomen; shining,
elytra without distinct reticukite micro-
sculpture. Head 0.75 and 0.76 width pro-
thorax; eyes separated from mouth below
by about 'c diameter of an vyv. Prothorax
transverse-subcordate; width/length 1 .43
and 1.45; base/apex 1.20 and 1.21; sides
converging and straight or slightly sinuate
before obtuse but well formed ( sometimes
slightly bhuited) basal angles; disc de-
pressed at sides near base, extensively punc-
tate, least so near middle; basal marginal
line faint or interrupted at middle. Klijlrtr.
width elytra prothorax 1.36 and 1.38; Stli
and 9th int(>rvals usually with a little sparse,
fine punctulation. Secotidanj sexual char-
acters normal. Measurements: length 8.3-
9.3; width 3.4-3.8 mm.
Ti/pcs. Ilolotvpe S (Bishop Mus.) and
3 paratypes (2 in M.C.Z., Type No. 31,372)
from Feramin, N-E. N. G., 1200-1500 m.
May 11-22 (holotype), 23-31, 1959 (W.
W. Brandt); and additional paratypes as
follows, all from N-E. N. G.: 3, Torricelli
Mts., Mobitei, 750 m. Mar. 16-31. Apr. 1-
15, 16-22, 1959 (W. W. Brandt, Bishop
Mus. ); 2, Eliptamin Vy., 1665-2530 m, June
23-30, and 2, same localitx, 1200-1350 m,
Aug. 16-30 and Sept. 1-15, 1959 (W. W.
Brandt, Bishop Mus.); 15, Wau, Morobe
Dist, 1100, 1200 (most), 1300, and 1700-
1800 m. Mar., Apr., and all months from
June to Dec., 1961-1963 (Sedlacek).
Additional material. The following addi-
tional specimens are tentatixely assigned
to this species. Papua: 1, Dogon, Amazon
Bay Dist., 2400 ft. (c. 730 m), Sept. 1962
(W. W. Brandt, C.S.I.R.O.); 1, Kokoda,
1200 ft. (366 m), July 1933 (Cheesman),
at light. N-E. N. G.: 1, Tuwep. Salawaket
Rge.\ 1350 m, Sept. 9, 1956 (E. J. Ford, Jr..
Bishop Mus.), in light trap. West N. G.:
1, Cyclops Mts., Sabron Camp 2. 2000 ft..
June 1936 (Cheesman); 1, "Neth. New
CTuinea' without further locality, Oct. 20.
1944 (T. Aarons, California Acad.).
Measured s})ecimens. The ^ holotype and
1 9 paratN'pe from Feramin.
Notes. See under lollowing species
(obscurus).
Trichotichnus obscurus n. sp.
Description. With characters of genus;
ionn broad-average; brownish piceous, lat-
eral margins of prothorax and cKtra \aguel\-
paler or translueeut; abdomen broadK' mar-
gin(>d with yellow, the \eIlow margins
widest anteriorh'; appendages testaceous,
hind leinora strikingK jiale; inodeiateK'
shining, elytia witli transNcrse reticulate
microsculpture \isible in both sexes (at
50 ■ ). Head 0.77 and 0.76 width prothorax;
eyes separated lioin mouth below by Vi
or less width o( an e\e. Prothorax trans-
The Carabid Beetles of New Guinea • Darlington 57
verse-subcordate; width length 1.40 and
1.44; base/apex 1.21 and 1.20; sides con-
verging, slightly, broadly sinuate before dis-
tinct but obtuse posterior angles; disc mod-
erately depressed at sides basally, extensively
but rather finely punctate except almost
impunctate at middle. Elytra: width
elytra/prothorax 1.34 and 1.27; 8th and 9th
intervals usually not distinctly punctulate.
Secondary sexual characters normal. Mea-
siiremeuts: length 8.5-9.4; width 3.3-3.7
mm.
Types. Holotype S (Bishop Mus.) and
2 paratypes (1 in M.C.Z., Type No. 31,373)
from Saidor, Matoko, Finisterre Rge., N-E.
N. G., Aug. 29-Sept. 5 and Sept. 6-24,
1958 (W. W. Brandt); and additional
paratypes as follows: Papua: 1, S. High-
lands^ Dimifa, SE. of Mt. Giluwe, 2200 m,
Oct. 11, 1958 (Gressitt). N-E. N. G.: 2,
Wau, Morobe Dist., 1200 m, Nov. 1-20 and
Dec. 1961 (Sedlacek); 7, Edie Creek, 14
km SW. of Wau, 2000 m, Feb. 13, 1962
(Sedlacek); 1, Eliptamin Vv., 1200-1350
m, Aug. 1-15, 1959 (W. W. Brandt, Bishop
Mus.).
Measured speciynens. The <^ holotype and
1 9 paratype from Saidor.
Notes. Although this species is superfi-
cially similar to the preceding one {hrandti),
I think it is distinct, differing most obvi-
ously by presence of elytral microsculpture.
The 8th and 9th elytral intervals are usually
less punctulate in ohscurus than in hrandti.
Jrichotichnus gutfula n. sp.
Description. With characters of genus;
form average; brownish black above, sides
of elytra ( and to some extent of prothorax )
testaceous, elytra with a conspicuous com-
mon testaceous subapical sutural spot
(reaching 3rd intervals), abdomen either
with irregular broad testaceous margins or
wholly rufescent, appendages brownish
testaceous; moderately shining, elytra usu-
ally with distinct transverse microreticula-
tion. Head 0.78 and 0.79 width prothorax;
eyes large, separated from mouth below
by c. Vs width of an eye. Prothorax sub-
transverse; width/length 1.48 and 1.43;
base/apex 1.19 and 1.21; sides rounded
anteriorly, converging and nearly straight
or slightly sinuate before obtuse but well
defined basal angles; disc depressed at
sides basally; basal marginal line usually
indistinct at middle; surface of disc exten-
sively punctate, the punctation finer and
less dense at middle. Elytra: width elytra/
prothorax 1.32 and 1.33; 8th and 9th inter-
vals not or not much punctulate. Secondary
sexual characters normal. Measurements:
length 7.6-8.7; width 3.1-3.6 mm.
Types. Holotype 6 (Bishop Mus.) and
31 paratvpes (some in M.C.Z., Type No.
31,374) 'from Wau, Morobe Dist., N-E.
N. G., 1200 m, Feb., Mar., June, Aug.,
Sept., Oct., Nov., Dec. 1961-1963 (Sedla-
ceks); 1 paratype, same locality, 1700-
1800 m, Nov. 17, 1961 (Sedlacek).
Additional material. Twenty-four speci-
mens from 13 widely scattered localities
including Dobodura, in all 3 political divi-
sions of New Guinea; altitudes, near sea
level to c. 2000 m. Specimens taken in
every month except May and June.
Measured specimens. The i holotype and
1 9 paratype from Wau.
Notes. This species is characterized by
its size plus presence of a conspicuous pale
subapical sutural spot. It is apparently
widely distributed especially in the foot-
hills and lower mountains of New Guinea.
Specimens of this or a closely related
species have been seen also from New
Britain and New Ireland (Bishop Mus.).
Trichotichnus mongi n. sp.
Description. With characters of genus;
form as in Figure 22, differing from other
Trichotichnus by subquadrate prothorax,
strongly narrowed anteriorly; piceous, lat-
eral margins of prothorax and elytra nar-
rowly translucent or pale, sutural intervals
reddish toward apex; shining, elytra sub-
iridescent, without distinct reticulate micro-
sculpture. Head 0.72 width prothorax;
eyes rather small, separated from mouth
below by more than V^ width of an eye.
58
Bulletin Museum of Comparative Zoology. Vol. 137, No. 1
Prothorox subquadrate except strongly nar-
rowed at extreme front; width length 1.40;
base/apex 1.35; sides rounded anteriorly,
broadly sinuate before c. right ( slightly
obtuse) basal angles; basal marginal line
fine but entire; disc scarcely depressed at
sides, extensively punctate, the punctures
finer and less dense across middle. Elytra
convex ( more so than usual ) ; width elytra/
prothorax 1.46; outer intervals with a little
sparse punctulation. Secondary .sexual char-
acters of 9 normal; i unknown. Measure-
ments: length c. 7.5; width 3.3 mm.
Type. Holotype 9 (M.C.Z., Type No.
31,375) from Tumnang, Mongi Watershed,
Huon Pen., N-E. N. G., 1400-1600 m, Apr.
14-15, 1955 (E. O. ^^^ilson); the type is
unique.
Notes. The unique form of this species
makes it worth describing, even though
I have only one specimen of it and do not
know the male.
Trichotichnus semirugosus n. sp.
Description. With characters of genus;
form c. average, small; brownish piceous,
margins of prothorax and elytra narrowly
testaceous, elytra with common subapical
sutural pale spot usually reaching 3rd in-
tervals; lower surface in part dark but with
extensive, irregular testaceous areas; anten-
nae brownish, legs brownish testaceous;
rather shining, elytra usually with faint
tran.sverse microreticulation. Head O.H\ and
0.84 width prothorax; eyes rather large,
separated from mouth below by <". 's width
of an eye. Prothorax subcordate; width
length 1.51 and 1.4(S; base apex 1.12 and
1.16; sides broadly round(>d anteriorly, con-
verging and straight or nearly so posteriorly;
posterior anglers distinct but obtuse, some-
times minutely denticulate; base more
oblique at sides than usual, not or indis-
tinctly margined; disc weakly depressed at
sides, extensively punctate, the punctation
finer and less dense across middle. Elytra:
width elytra /prothorax 1.39 and 1.44; 8tli
and 9th intervals rugosely punctate, punc-
tation present but less dense at bases of 6th
and 7th intervals. Secondary sexual char-
acters normal. Measurements: length 5.3-
6.8; width 2.3-2.8 mm.
Types. Holotype 9 (M.C.Z., Type No.
31,376) and 2 (99) paratypes from
Dobodura, Papua, Mar.-July 1944 (Dar-
lington); and additional paratvpes as fol-
lows: N-E. N. G.: 1, Wau, 1200 m, No\'.
21, 1961 (Sedlaceks); 7, Finschhafen, Huon
Pen., 10 m, Apr. 9-16, 1963 (Sedlacek), in
mercurv vapor light trap; 3. Torricelli Mts.,
Mobitei, 750 m, Feb. 28-Mar. 4, Mar. 16-
31, 1959 (W. W. Brandt, Bishop Mus.).
West N. G.: 1, Cyclops Mts. (no further
details) (Cheesman); 2, Hollandia area.
W. Sentani, Cyclops Mts., 50-100 m, June
22-24, 1959 (Gressitt), in light trap; 2,
Ifar, Cvclops Mts., 450-500 m, Sept. 7 and
9, 1962 (Sedlacek).
Measured specimens. One i paratype
from Mobitei and the 9 holotype, in this
order.
Notes. The coloration and the dense
punctation of the ehtral margins would
place this species in Lampetes, if Lampetes
were distinguished from Trichotichnus. The
new species is in fact close to Lampetes
isahellinus Louwerens of Amboina (Tijd-
schrift voor Ent. 105, 1962, p. 140). Ho\\'-
ever, comparison with paratopes of isa-
hellinus shows that scmiruii.osus has the
outer elytral intervals more completeK' and
more densely rugose, although the differ-
ence is not great.
Trichotichnus mixtus n. sp.
Descriplion. With characters of genus;
lorm average, rather small; brownish i^ice-
ous, lateral margins ol prothorax and elytia
transluc(Mit or pale, and sutural and some-
times 2nd interxals ol clxtia paler befori'
apex (variable), abdomen broadK" but ir-
regularly pale-margined, legs and antennae
irregularly brownish ti'staceous; shining,
elytra with or without light transverse
mieroretieulation. llccul 0.79 and 0.79 width
piothorax; eyes rather large, separated Irom
mouth below by c. 'is diameter ol an eye. Pro-
ihonix transNcrse-subeordate; width length
The Carabid Beetles of New Guinea • Darlington 59
1.54 and 1.54; base apex 1.15 and 1.13;
sides broadly rounded anteriorly, nearly
straight and converging posteriorly to ob-
tuse, sometimes slightly blunted posterior
angles; basal marginal line usually inter-
rupted at middle; disc depressed at sides
basally, extensively punctate, the puncta-
tion finer and sparser across middle. Elytra:
width elytra prothorax 1.36 and 1.36; 7th
and 8th intervals variabK' punctate (8th
varying from almost impunctate to almost
rugose); other intervals sparsely or not
punctulate. Secondary sexual characters
normal. Measurements: length 5.8-7.5;
width 2.8-3.2 mm.
Types. Holotype S (Bishop Mns.) and
4 paratypes (2 in M.C.Z., Type No. 31,377)
from Torricelli Mts., Mobitei, N-E. N. G.,
750 m. Mar. 5-15, Apr. 16-22 (holotype
with latter date), 1959 (W. W. Brandt).
Additional material. Twenty-one speci-
mens from 11 localities (including Wau)
in all 3 political divisions of New Guinea
are assigned to mixtus but not as types.
They vary considerably in several char-
acters.
Measured specimens. The c5 holotype and
1 9 paratype.
Notes. Because of the variation of this
species ( if it is all one species ) I have con-
fined the type series to specimens from one
locality. In general, the species should be
recognizable by size; sutural intervals pale
before apex; prothorax usually relatively
wide (wider and a little more depressed
at sides toward base than in scmirug,o.sus);
and outer elytral intervals usually punc-
tulate but not rugose, although this last
character is surprisingly variable even in
the type series.
Trichotichnus delicafus n .sp.
Description. With characters of genus;
form ( Fig. 23 ) slender-average, very small;
brownish piceous, prothorax with margins
narrowly pale or translucent, elytra with
margins and common subapical sutural spot
(including small parts of 2nd intervals)
testaceous or rufescent; abdomen with or
without well defined pale margins; af)pend-
ages testaceous; shining, elytra without
reticulate microsculpture. Head 0.74 and
0.77 width prothorax; eyes moderate, sepa-
rated from mouth below by c. '^i\ or Vs width
of an eye. Prothorax subcordate; width/
length 1.50 and 1.44; base/apex 1.11 and
1.21; sides broadly slightly sinuate before
obtuse, usually slightly blunted posterior
angles; basal marginal line vague or incom-
plete at middle; disc scarcely depressed at
sides even basally, less densely punctate
than usual, with middle of disc least punc-
tate. Elytra: width elytra/ prothorax 1.30
and 1.31; outer intervals without or with
only sparse punctulation. Secondary sexual
characters normal. Measurements: length
5.3-5.8; width 2.2-2.4 mm.
Types. Holotype 9 (Hungarian Nat.
Mus.) and 1 9 paratype (M.C.Z., Type
No. 31,378) from I. Deslacs (Garove Is.),
N-E. N. G., 1901 (Biro); and additional
paratypes as follows: Papua: 1, Woodlark
Is. (Murua), Kulumadau Hill, Mar. 9-12,
1957 (W. W. Brandt, Bishop Mus.). West
N. G.: 1, Hollandia, Dec. 1944 (W. T.
Nailon, Fenton Coll.); 1, Res. Hollandia,
Dojo, 2nd Strip, July 12, 1957 ( R. T. Simon
Thomas, Louwerens Coll.).
Measured specimens. A i paratype from
Woodlark Is. (the only 6 of the species
seen) and the 9 holotype, in this order.
Notes. T. delicatus is characterized b\'
small size, markings, and scarcely depressed
sides of pronotum.
Genus HARPALOXENUS Schauberger
Schauberger 1933, Ent. Anzeiger 13, p. 154.
Diagnosis. Characters as for Trichotichnus
(preceding genus) except anterior tibiae
wider; form characteristic (Fig. 24); upper
surface without distinct reticulate micro-
sculpture, but 8th and 9th elytral intervals
closely punctulate (except in fortis), other
intervals sparselv or not punctulate; wings
full.
Description. None required here.
60
Bulletin Museum of Cotnporaiive Zoology, Vol. 137, No. 1
Type species. 11. javanus Schauberger, of
Java.
Generic (listriJ)iifion. Java and Ando-
nare Is., Celel>es, Philippines, Moluccas
(Halmahera), New Guinea, and (unpub-
lished) Solomon Islands and New
Hebrides.
Notes. The species assigned to this genus
seem to form a natural group. However,
the group is apparently closely allied to
TricJioticlinus and further study may show
that it is not worth generic separation.
Key to Species of Harpaloxenus of
New Guinea
1. Anterior tibiae with apex c. Mi wide as
til)ial leii.irtli: head relatively wider (more
than 0.80 width prothorax) - ^ - 2
- Anterior tibiae with apex c. M wide as tibial
len.qth; head relatively smaller (usually less
than 0.80 width prothorax) -- 3
2. Male with front but not middle tarsi
squamulose (p. 60) fortis
- Male with front and middle tarsi squamulose
(p. 61) ma.s
3. Larj^er (length c. 11 mm) (p. 62) wan
- Smaller (length c. 10 mm or less) - 4
4. Frouotum with sides usually slightly de-
pressed toward 1)ase, base usually more
eoarsely and extensiveh' punetate, posterior
angles usually better defined, and median-
lateral setae usually c. Vi of prothoracic
length before apex (p. 61) celehen.sis
- Pronotum with sides not depressed, base
usualK' more finely punetate especially at
middle of base, posterior angles usually more
obtuse, and median-lateral setae usually c.
'':! of prothoracic length before apex (p.
62 ) sedlaccki
Harpaloxenus fortis n. sp.
Dcscri])lio)i. With characters of genus;
form as in Figure 24, heavily built; ])rownish
piceous, lateral margins ol pronotum and
elytra including 9th intervals ±: yellowish,
lower surhice with extensive yellowish areas
especially laterally, appendages brownisli
testaceous. Head wide, ().<S(S and ().(S5 w idth
prothorax; antennae stout, middle segments
scarcely longer than wide. Proihorax
broadly cordate; width/length 1.57 and
1.58; base/apex 0.93 and 0.99; sides con-
verging and usually l)roadl\' but not strongly
sinuate before obtuse but distincl basal
angles; disc slightly depressed at sides pos-
teriorly, basal impressions weak and ir-
regular, base punctate chiefly toward sides.
Elytra elongate-quadrate; width elytra pro-
thorax 1.14 and 1.14; outer intervals (8.
9) less punctulate than usual in genus; 3rd
intervals either with or (usually) without
minute puncture on inner edge behind
middle. Le^.s: front tibiae very wide in
both sexes, apex c. Va wide as tibial length,
and apex usually sinuate-emarginate with
outer angle slightly produced; middle tibiae
slightly wider and more arcuate than in
most other species of genus. Secondary
sexual characters: 6 front tarsi slightly di-
lated, with segments 1 (apex only) to 4 2-
seriately squamulose below; middle tarsi
not perceptibly dilated and not squamulose.
Measurements: length 8.5-10.3; width 3.1-
3.7 mm.
Types. Holotype S (M.C.Z., Type No.
31,379 ) and 17 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes from Papua as fol-
lows: 8, Kokoda, 1200 ft. (366 m). May.
July, Aug. 1933 (Cheesman); 1, same lo-
cality. Mar. 20, 1956 (Gressitt), in light
trap; 2, Saputa, near Buna, 1943-44 ( R. B.
Speiry, Chicago Mus.); 1, Deria, Amazon
Bay Dist., Dec. 1962 (W. W. Brandt.
C.S.I.R.O.); 1, Mt. Lamington, 1300-1500
ft. (c. 400-450 m) (C. T. McNamara, S.
Australian Mus.).
Additional material. West N. G.: 2.
(99), Ilollandia, Apr. 1945 (B. Malkin.
U.S.N.M.); 3 (99), Hollandia area, W.
Sentani, Cyclops Mts., 50-100, 150-250 m,
June 17, 22-24, 1959 (Gressitt and T. C.
Maa, Bishop Mus.), 2 of thes(> specimens
teneral and taken in light trap; 1 s" , Kota
Nika, Res. Ilollandia, Jan. 25, 1956 ( R. T.
Simon Thomas, Louwerens (^oU.); 1 9,
Wasian, Vogelkop, Sept. 1939 (Wind.
M.C'.Z. ). Thes(> specimens unfortunat<:4\
are all 9 9 . The\' arc referred to the
present spc>cies rather than the following
one (mas) because the 8th and 9th ehtral
inter\als are almost impunctate.
The Carabid Beetles of New Guinea • Darlington 61
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. The present species differs from
all others known in the genus by absence
of squamules on the male middle tarsi.
However, I do not think that this justifies
making a separate genus or subgenus. This
species seems otherwise to be a well char-
acterized Harpaloxenus, and it apparently
is closely related to the following (mas),
which has the male middle tarsi normally
squamulose.
Harpaloxenus mas n. sp.
Description. With characters of genus;
form c. as in preceding species (foiiis)
but slightly less heavily built; characters as
in preceding species except as follows.
Head relatively slightly smaller, 0.81 and
0.S2 width prothorax. Prothorax with sides
converging but not or scarcely sinuate pos-
teriorly, and with posterior angles slightly
more obtuse; width/length 1.52 and 1.45;
base/apex 1.00 and 0.9cS. Elytra: width
elytra/prothorax 1.24 and 1.29; outer inter-
vals (8, 9) closely punctulate at least an-
teriorly, sometimes in part rugose. Sec-
ondary sexual characters: i front and middle
tarsi slightly dilated, 2-seriately squamu-
lose. Measurements: length c. 9.5-10.5;
width 3.5-3.7 mm.
Types. Holotype $ (Bishop Mus.) and
3 ( (5 9 9 ) paratypes (pair in M.C.Z., Type
No. 31,380) from Finschhafen, Huon Pen.,
N-E. N. C, 10 m, Apr. 9-16, 1963 ( Sedla-
cek), in mercury vapor light trap; 1 i
paratype, Wau, 1050 m, Nov. 4, 1961
(Sedlacek); 1 i paratype. Hoi Maffin, near
Sarmi, West N. G., July 18, 1959 (T. C.
Maa, Bishop Mus.).
Measured specimens. The 6 holotype and
1 9 paratype from Finschhafen.
Notes. This species resembles the pre-
ceding one in width of front tibiae but
differs in details of form (slightly nar-
rower head, slightly differently .shaped pro-
thorax) and in extensive punctulation of
outer elytral intervals. The paratype from
Hoi Maffin has these intervals less punc-
tulate than in the Finschhafen specimens
but still more punctulate than in any fortis
that I have seen.
Harpaloxenus celebensis Schauberger
Schauberger 1933, Ent. Anzeiger 13, pp. 155, 157.
Louwerens 1953, Verhandlungen Naturforschenden
Gesellschaft Basel 64, p. 306.
1956, Treubia 23, p. 222.
Description ( for recognition only ) . Form
average, rather variable; prothorax and
elytra usually narrowly yellow-margined.
Head 0.75 and 0.81 width prothorax. Pro-
thorax trans verse-subcordate; width length
1.50 and 1.52; base/apex 1.20 and 1.06 (ex-
ceptionally variable); sides rather weakly
converging, not or only slightly sinuate be-
fore distinct but slightly obtuse or blunted
posterior angles, with anterior-lateral setae
usually c. V4 of prothoracic length from
apex. Elytra: width elytra/prothorax 1.24
and 1.21; dorsal punctures usually present,
on inner edge 3rd intervals against 2nd
striae (sometimes absent on one or both
elytra). Legs: front tibiae with apex c. %
wide as tibial length. Secondary sexual
characters: S front and middle tarsi squamu-
lose. Measurements: length 8.5-10; width
3.2-3.8 mm.
Type. From South Celebes; probably
in Schauberger Coll. (not seen).
Occurrence in New Guinea. Papua: 7,
Dobodura, Mar.-July 1944 (Darlington);
2, Dogon, Amazon Bay Dist., 2400 ft. (c.
740 m), Oct.-Nov. 1962 (W. W. Brandt,
C.S.I.R.O.) N-E. N. G.: 3, Wau, Morobe
Dist, 1200 m, Dec. 18, 1961 (Sedlacek);
1, Finschhafen, 10 m, Apr. 9-16, 1963
(Sedlacek), in hght trap; 2, Torricelli Mts.,
Mobitei, 750 m. Mar. 5-15, 16-31, 1959
{\y. W. Brandt, Bishop Mus.). West N. G.:
6, Hollandia and vicinity including Cyclops
Mts. (various dates and collectors); and 5,
doubtfully identified, from localities farther
west in West N. C, including Biak Is.
Measured specimens. A pair ( <^ 9 ) from
Dobodura.
Notes. Louwerens records celebensis
from Java, Sumba, and Halmahera, as
62 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
well as Celebes, and the present records
extend its range to New Guinea. However,
variation is considerable and I am not sure
of my identifications in some cases. Small
specimens of this species can be confused
with certain Trichotichnus, especially with
discolored individuals of ii.uttula, but the
particular Trichotichnus in question do not
have the Sth and 9th elvtral intervals closelv
punctulate and have, of course, slightly
narrower front tibiae. See under the follow-
ing species (sedlaceki) for further com-
parisons.
Harpaloxenus sedlaceki n. sp.
Description. With characters of genus;
form average; black or piceous, sides of pro-
notum not or vaguely pale, elytra narrowly
pale-margined, abdomen pale-spotted at
sides, appendages brownish testaceous.
Head 0.75 and 0.75 width prothorax.
Prothorax transverse-subquadrate; width
length 1.50 and 1.49; base apex 1.19 and
1.21; sides moderately converging and nearly
straight posteriorly but usually not sinuate,
with median-lateral setae usually c. M', of pro-
thoracic length before apex; posterior angles
obtuse, ± blunted; disc rather strongly almost
evenly convex, not depressed at sides pos-
teriorly, baso-lateral impressions slight and
poorly defined; base (rather finely) punctate
especially at sides. Elytra: width elytra, pro-
thorax 1.22 and 1.20; outer intervals (8, 9)
extensively closely punctulate; dorsal punc-
tures of 3rd intervals usually c. midway be-
tween 2nd and .3rd striae (see Notes, be-
low). Le^s: front tibiae with apex c. Vl
wide as tibial length. Secondary sexual
characters: 6 front and middle tarsi sliglith
dilated, 2-seriately scpiamulose. Mea.surc-
menls: l(>ngth 8.0-9.0; width 3.1-3.5 mm.
'Types. Holotype $ (Bishop Mus.) ami
20 paratypes (some in M.C.Z., Type No.
31,381) from Wau, Morobe Dist., 'l20() m,
N-E. N. G., dates in Apr., Aug., Oct., Nov.,
Dec. 1961-1963 (Sedlacek).
Additional material. Twenty-two speci-
mens from numerous localities, from Mo-
dewa and Dobodura in Papua to Hollandia
in West N. G. Some of these specimens
are identified only doubtfully.
Measured s))ecimens. The i holotype and
1 9 paratype.
Notes. This species is similar to celehensis
(above) but differs in having the pronotum
slightly more convex with sides usually not
at all depressed toward base, and in other
ways indicated in the preceding Key to
Species of Harpaloxenus of Neic Guinea.
In the types of sedlaceki the dorsal punc-
ture of the 3rd interval is usually midway
between the 2nd and 3rd striae, not close
to the 2nd stria. Of the 21 specimens from
Wau, only 1 has the puncture close to the
2nd stria (on the inner edge of the 3rd
interval) on both elytra. Two have the
puncture close to the 2nd stria on one
elytron but near the middle of the 3rd
interval on the other. One has the puncture
close to the 3rd stria on one elytron. And
17 have the puncture near the middle of the
3rd interval (but somewhat variable in
position ) on both elytra. However, this un-
usual position of dorsal punctures may be
characteristic of the local population at
Wau rather than of the species as a \\'hole.
Harpaloxenus wau n. sp.
Description. With characters of genus;
form nearly as in celehensis and sedlaceki,
but larger; pronotum and ehtra not or nar-
rowly and faintly pale-margined, abdomen
with irregular testaceous marks at sides,
appendages brownish testaceous. Head 0.73
and 0.75 width prothorax. Prothorax trans-
verse-subquadrate; width length 1.48 and
1.51; base apex 1.17 and 1.24; sides slightly
converging posteriori), nearly straight but
not sinuate before slightK obtuse-blunted
posterior angl{\s; disc usualK' slightly de-
pressed at sides posteriorh', baso-lateral im-
pressions \agu(\ base finch punctate espe-
cialh- toward sides. Elytra: width elytra
prothorax 1.30 and 1.28; outer intervals (8,
9) extensively punctulat(\ 3rd intervals with
dorsal punctur(> usualK' by 2nd stria behind
middle. Leii,s: Iront tibiae with ajiex c. 'i
wide as tibial length. Secondary sexual
The Carabid Beetles of New Guinea • Darlington
63
characters: S front and middle tarsi slightly
dilated, 2-seriately squamulose. Measure-
ments: length c. 11; width c. 4.2 mm.
Types. Holotype c^ (Bishop Mus.) and
29 paratypes (some in M.C.Z., Type No.
31,382) from Wau and vicinity, Morobe
Dist., N-E. N. G., 1100, 1200 (most), 1700-
1800 m, dates in every month, 1961-1964
( Sedlacek ) .
Measured specimens. The 6 holotype and
1 9 paratype.
Notes. The large size distinguishes this
species from other similar ones. As com-
pared with sedlaceki, with which it occurs,
wau is not only larger but has the sides of
the pronotum usually slightly depressed
toward base and the dorsal puncture of
the 3rd intervals usually adjacent to the 2nd
stria. Of the 8 types, only 1 has this punc-
ture distant from the 2nd stria on both
elytra. Two have the puncture against the
2nd stria on one elytron but distant from it
on the other elytron. And 5 have the punc-
ture against or very near the 2nd stria on
both elytra.
LYTER n. gen.
Diapiosis. Form and characters of me-
dium-sized Trichofichnus, but c$ front and
middle tarsi below with more than 2 rows
of long, slender scales forming a loose
vestiture (not a dense sole); prosternum
glabrous anteriorly; 3rd elytral intervals
without or with only faint traces of dorsal
punctures.
Description. Head: eyes separated from
mouth below by c. Vs diameter of an eye;
antennae rather short, middle segments c.
IVaX long as wide; front smoothly convex
except frontal suture shaiply impressed,
with impressed lines extending diagonally
back to above eyes; mentum with triangular
tooth; labial palpi with penultimate seg-
ments plurisetose; ligula long, emarginate,
2-setose outside middle of length; para-
glossae shorter than ligula. Frothorax with
1 lateral seta each side about Vi prothoracic
length from apex. Elytra: margins entire
at base, obtusely subangulate at humeri,
sinuate before apex; marginal channels nar-
row, without partial 10th intei-vals; striae
entire; scutellar striae long, at base 2nd
intervals; 3rd intervals impunctate or with
vestige of puncture by 2nd striae behind
middle (position as in Trichotichnus) .
Lower surface: prosternum glabrous an-
teriorly but with several setae at apex pro-
sternal process. Inner wings full. Legs:
front tibiae irregularly truncate, apex c. V:,
wide as tibial length, with principal (inner
apical) spur not much expanded; hind tarsi
with 1st segment 2x or more as long as wide
at apex, 5th segment with 2 accessory setae
each side. Secondary sexual characters: S
front tarsi slightly dilated, 4 segments
loosely clothed below with slender long
scales; middle tarsi scarcely dilated, with
some (fewer) similar scales; 2 setae each
side last ventral segment in both sexes;
S copulatory organs as in Figure 174, with
apex of middle lobe not produced beyond
orifice.
Type species. Lyter glaher n. sp. (below).
Generic distribution. The single species
is confined to New Guinea, so far as
known.
Notes. The relationships of this new
genus are doubtful. It may be derived
from an ancestor like Trichotichnus medius
(p. 55), from which it differs most obvi-
ously in the clothing of the 6 tarsi (2-
seriately squamulose in Trichotichnus). The
new genus is notable also for its relatively
long ligula, for absence of pubescence on
anterior part of prosternum, and for virtual
suppression of dorsal elytral punctures.
The name Lyter, from the Greek, signifies
one who loosens ( the squamae of the S front
tarsi ) .
Lyter glaber n. sp.
Description. With characters of genus;
form as in Figure 25; reddish piceous, ap-
pendages redder; shining, reticulate micro-
sculpture absent on front of head and disc
of pronotum, faint and somewhat transverse
on elytral intervals. Head 0.75 and 0.77
width prothorax. Prothorax: width length
64 Bulletin Museum of Comparative Zoolofiy. Vol. 137, No. 1
1.45 and 1.49; base/apex 1.12 and 1.16;
sides rounded anteriorly, c. straight and
converging in posterior half, narrowly mar-
gined; posterior angles slightly obtuse,
blunted; base and apex with or without
faint marginal lines; disc slightly depressed
at sides basally, the depressed areas finely
but not closely punctate. Elytra: width
elytra prothorax 1.21 and 1.22. Measure-
ments: length 7.3-8.3; width 2.9-3.3 mm.
Types. Holotype $ (California Acad.)
and 32 paratypes (some in M.C.Z., Type
No. 31,383) from Finschhafen, N-E. N. G.,
various dates in April and May (holotype.
May 1) (E. S. Ross); and additional para-
types as follows. Papua: 21, Kokoda,
1200 ft. (366 m), Apr., Aug., Sept. 1933
(Cheesman); 1, Owen Stanley Rge., Goilala,
Tapini, 975 m, Nov. 16-25, 1957 (W. \V.
Brandt, Bishop Mus.); 1, Dogon, Amazon
Bay Dist., 2400 ft. (730 m), Oct.-Nov.
1962 (W. \V. Brandt, C.S.I.R.O.); 1, Mt.
Lamington, 1300-1500 ft. {c. 400-450 m)
(C. T. McNamara, S. Australian Mus.).
N-E. N. G.: 5, "No. 14," Umi R., Markham
Vy., 480 m, dates in Nov. 1959 (Sixth
Archbold Exp., A.M.N.H.); 1, Lae, July
1924 (F. E. Skinner, Bishop Mus.); 3,
Bulolo, 730, 1170 m, Aug. 15, 19, 21, 1956
(E. J. Ford, Jr., Bishop Mus.); 58, Wau,
Morobe Dist., 1200 m, dates in every
month except June, 1961-1963 (Sedlacek);
2, same locality, 1700-1800 m, Nov. 17,
1961 (Sedlacek); 4, Sum-Sum, 64 km N. of
Wau, 580 m, Feb. 15, 1963 ( Sedlacek ) ; 1,
Karimui, S. of Ooroka, 1000 m, June 3,
1961 (Cre.ssitt), in light trap. West N. <;.:
3, Ilollandia area, \V. Sentani, Cvclops
Mts., 50-100, 150-250 m, June 17, 22-24,
1959 (Gressitt); 1, Kota Nika, Res. Ilol-
landia, Feb. 14, 1956 (R. T. Simon Thomas,
Louwerens Coll.), in light trap.
Measured speeimens. The i holotype and
1 9 paratype Irom I'^inschhalen.
Notes. Although this carabid is appar-
ently common in some places, I hiiled to
find it and do not know its habitat. The
k)calities suggest that it lives in rain forest.
Specimens Irom Bulolo and Ilollandia were
taken in light traps, which implies that the
insect flies.
Genus COLEOUSSUS Bates
Bates 1892, Ann. Mus. Civ. Genoa 32, p. 338.
Csiki 1932, Coleop. Cat., Caral)idae, Harpalinae 6,
p. 1217 (as suligenus of TriclioticJiniis) .
Andrewes 1939, Ann. Maj?. Nat. Hist. (11) 3, p.
132.
Diaiinosis. See Key to Genera of Har))alini
of New Guinea.
Deseription ( for recognition only ) . Form
of broad medium-sized Harpalini; upper
surface ( in New Guinean species ) \'er)-
shining, without reticulate microsculpture.
Head: eyes relatively large (compared
with most Triehotichnus), almost contigu-
ous with sides of mouth below; frontal
impressions deep, subpunctiform or curving
toward eyes posteriorly; mentum toothed;
ligula rounded, 2-setose; paraglossae at-
tached to ligula but longer, with narrowly
rounded apices; penultimate segments labial
palpi with more than 2 setae anteriorh'.
Prothorax as in Trieliotiehnus. Elytra:
sutural angles denticulate ( in New Ciuinean
species); striae entire; sutural striae long;
3rd intervals seriate-punctate. Inner winiis
full. LeiS,s: tarsi slender. Secondary .sexual
characters: see Descriptions of species.
Ty))e species. llypolitJius perlueens Bates,
of Kashmir, etc. (fixed In- Andrewes, 1939).
Generic distribution. India and Ceylon,
Sikkini, Burma, etc., to Java, Borneo,
Philippines (Negros), Cele]>es, Burn,
Moliiecas (Amboina), New (Guinea, Solo-
mons, New Hebrides, and ihc Cape York
Pen. of Australia (occurrence in Philii")-
pines. New Hebrides, and Australia bas(>d
on unpublished records ) .
Notes. Members of this genus seem to
be rare insects, usualK' taken onl\ one or
two indi\iduals at a time, although the\
are winged and IK to light. I do not know
their habitat.
Key io Species of Coleolisscs of
Ni:\\ (".riNKA
I. Onter an,ul(\s of cKtia not (Iclincd. hroaill)
ronndecl (p. 65) jHijitui
The Carabid Beetles of New Guinea
Darlington
65
- Outer angles of elytra ( before subapical
sinuations) well defined, right or obtuse (p.
65 ) angulatns
Coleolissus papua n, sp.
Description. With characters of genus;
form as in Figure 26, large, broad; lilack,
appendages irregularly reddish piceous;
very shining, elytra iridescent. Head 0.73
and 0.72 width prothorax; front faintly
punctulate. Prothorax transverse; width/
length 1.54 and 1.56; base/apex c. 1.11 and
1.11; sides broadly rounded to rounded
posterior angles; lateral margins broader
and more depressed posteriorly; apex mar-
gined, base indistinctly so; disc depressed;
baso-lateral impressions broad but poorly
defined, closely punctate; other parts of
disc sparsely or not punctulate. Elytra
wide; width elytra prothorax 1.30 and 1.39;
base margined; humeri rounded; outer sub-
apical angles rounded; apices slightly sinu-
ate before denticulate sutural angles; striae
impunctate; intervals slightly convex, finely
sparsely (scarcely detectably) punctulate,
3rd with c. 7 small punctures irregularly
spaced along most of length of inner edge.
Lower siwface: prosternum and abdomen
with a little fine, short, sparse pubescence
(scarcely detectable); prosternal process
setose. Secondary sexual characters: S un-
known; 9 with 2 setae each side last
ventral segment. Measurements: length
12.5-13.5; width 4.8-5.3 mm.
Types. Holotype 9 (Bishop Mus.) from
Kiunga, Fly R., Papua (W. W. Brandt);
and 1 9 paratype (M.C.Z., Type No.
31,384) from Hollandia area, VV. Sentani,
Cyclops Mts., West N. G., 150-250 m, June
23, 1959 (T. C. Maa).
Notes. This species is close to Coleolissus
leveri Van Emden of the Solomons ( I have
a specimen compared with the type) and
even closer to C. kalisi Louwerens of Celebes
( I have 2 paratypes received by courtesy of
Mr. Louwerens). The New Guinean insect
is slightly larger and broader than kalisi,
with broader baso-lateral prothoracic im-
pressions. I collected a single specimen of a
Coleolissus very similar to pai)ua on the
Cape York Pen., Australia, in 1958.
Coleolissus angulatus n. sp.
Description. With characters of genus;
form as in Figure 27; black, elytra subirides-
cent, appendages reddish brown. Head 0.72
and 0.72 width prothorax; front virtually
impunctate. Prothorax transverse-subcor-
date; width/length 1.51 and 1.56; base/apex
1.12 and 1.09; sides converging but usually
not sinuate (sometimes slightly so) before
± rounded posterior angles; lateral margins
moderately broad and reflexed; apex mar-
gined, base not; disc weakly convex, baso-
lateral impressions not sharply defined,
rather closely but irregularly punctate; disc
otherwise less closely but extensively punc-
tate especially across base and at sides,
scarcely punctate at middle. Elytra: width
elytra prothorax 1.30 and 1.32; basal mar-
gin entire, vaguely obtusely angulate at
humeri; outer apical angles well defined,
usually right, sometimes obtuse; sutural
angles denticulate; striae deep, impunctate;
intervals slightly irregular but not distinctly
punctate, 3rd with c. 7 very small punctures
irregularly spaced on inner edge along
most of length of intei^vals. Lower suiiace:
prosternum not pubescent except for setae
at apex of prosternal process; abdomen not
pubescent except for usual "fixed" setae.
Secondary sexual characters: $ front tarsi
moderately dilated, middle tarsi scarcely
so, both pairs 2-seriately squamulose; 2
setae each side last ventral segment in
both sexes. Measurements: length c. 7.5-
8.5; width 3.1-3.3 mm.
Types. Holotype S (M.C.Z., Type No.
31,385) and 2 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. N-E. N. G.:
3, Finschhafen, 10 and 180 m, Apr. 9-16,
1963 (Sedlacek); 1, Aitape, Aug. 1944
(Darlington). West N. G.: 2, Hollandia
area, W. Sentani, Cyclops Mts., 150-250 m,
June 25, 1959 (Gressitt and T. C. Maa,
Bishop Mus.); 1, Ifar, Cyclops Mts., 150-
66
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
500 m, Sept. 6-9, 1962 (Sedlacek); 1, "Neth.
New Guinea" without further loeality, Dec.
10, 1944 (T. Aarons, California Acad.).
Measured specimens. The <:5 holotype and
1 9 paratype from Dobodura.
Notes. The deep elytral striation and
sharply defined outer elytral angles well
characterize this Coleolissus. The Dobodura
specimens were, I think, taken at light, and
some specimens from other localities are
evidently from light trap material.
Genus HYPHAEREON Macleay
Macleay 1825, Annulosa Javanica, p. 22.
Andrewes 1919, Trans. Ent. Soc. London for
1919, p. 156.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1221 (see for additional references).
Diapiosis. Among New Guinean Har-
palina this genus is recognizable by: form
rather NebrioAike; elytra with series of
(small) punctures on inner edge 3rd inter-
vals; only 1 seta each side last ventral seg-
ment in both sexes.
Description ( for recognition only ) . Form
c. Nehria-like, convex; reticulate micro-
sculpture faint or absent on head and
pronotum, fine and transverse ( ± visible at
50x ) on elytra; elytra subiridescent. Head:
eyes moderate, narrowly separated from
mouth below; mentum toothed; ligula sub-
truncate, 2-setose; paraglossae rounded, at-
tached to but longer than ligula. Frothorax
subcordate; anterior marginal line fine but
usually entire. Elytra with margins sinuate
before apex; striae entire; 3rd intervals
seriate-punctate. Inner uini!,s: see under
species. Secondary sexual characters: i
front and middle tarsi slightly dilated, 2-
seriately squamulose; 1 seta each side apex
last ventral segment in both sex(\s.
Type species. II. reflexus Macleay, of
Java.
Generic distrihtition. Known from Su-
matra, Java, Flores, Celebes, and New
(riiinea (not Australia).
Notes. Of the 3 New Guinean species of
Ilyphaercon, timid us is most like the type
species, with whicli I shall compare it (in
Notes under timidus). Calathomijnus, which
resembles Ilyphaercon in form and in having
only 1 pair of setae on the last ventral
segment in both sexes, and which also oc-
curs in the Malay Archipelago (but not
New Guinea ) , is probably closely related
to Hyphacreon but differs in having strongly
angulate humeri.
Key to Species of Hyphaereon of
New Guinea
1. Humeral margins broadly evenly rounded;
elytral striae shallow, inter\'als flat; setae of
apical ventral segment distant from margin
by more than ^/io length of segment (p.
66 ) levis
- Humeral margins obtusely subangulate; elytral
striae deeper; setae of last ventral segment
less than %o length of segment from margin - 2
2. Prothorax less cordate, with slightly broader
base (width of base/width of head 1.20 and
1.17); wings dimorphic, often much reduced;
lowland-living (p. 67) timidus
- Prothorax more cordate, with narrower base
(width of base of prothorax/ width of head
1.14 and 1.12); wings large, folded; moun-
tain-living (p. 68) cordcHfi
Hyphaereon levis n. sp.
Description. With characters of genus;
fonn ( Fig. 28 ) average; black, legs testa-
ceous, antennae and mouthparts brown.
Head 0.67 and 0.66 width prothorax.
Prothorax subcord;ite-subquadrate; wddth
length 1.38 and 1.38; base/apex 1.19 and
1.16; base/head 1.10 and 1.07; sides con-
verging and very slightly sinuate before
obtuse, blunted posterior angles; baso-kit-
eral impressions poorh' defined; disc ex-
tensively pimctate especially across base
and at apex, almost impunctate across mid-
dle. Elytra: width elytra prothorax 1.26
and 1.26; details ;is usu;il in genus except
humeral margins broadly evenly rounded;
striae entire^ but less deep than usual, hmer
u'inis,s lull in both specimens. Measure-
ments: l(Migth c. 9.0; ^^■idth 3.3-3.4 mm.
Types. Ilolotvpe $ (Leiden Mus. ) and
1 5 "paratype (M.C.Z., Type No. 31,386)
both from Sibil, Star Rge., West N. G..
1260 in, |une 1959 (Neth. New Gnincvi
Kxp.).
The Carabid Beetles of New Guinea • Darlington
67
Notes. For comparisons, see preceding
Key.
Hyphaereon timidus n. sp.
Description. With characters of genus;
form (Fig. 29) of small, rather broad
Nebria; black or piceous, appendages testa-
ceous, antennae in part brown. Head 0.64
and 0.65 width prothorax. Prothorax sub-
cordate-subquadrate; width /length 1.42 and
1.39; base/apex 1.20 and 1.17; base/head
1.20 and 1.17; sides converging and usually
slightly, broadly sinuate before obtuse,
slightly blunted posterior angles; anterior
marginal line entire or not (variation in-
dividual); baso-lateral impressions poorly
defined; disc finely irregularly punctate
basally, c. impunctate elsewhere. Elytra:
width elytra prothorax 1.19 and 1.23;
humeri obtusely but usually distinctly angu-
late; striae deep, intervals convex. Wings
dimorphic or polymorphic (see Notes, be-
low). Secondary sexual characters as for
genus. Measurements: length c. 6-7;
width 2.6-2.9 mm.
Types. Holotype i (M.C.Z., Type No.
31,387) and 55 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); 20
paratvpes from Oro Bay (near Dobodura),
Dec. 'l943-Jan. 1944 (Darlington).
Additional material. N-E. N. G.: 17,
Nadzab, July 1944 (Darhngton); 1, same
locahty, June 1944 (Krombein, U.S.N.M.);
1, Erima, Astrolabe Bay, 1896 (Biro); 1,
Busu R., "12 km," Sept. 21, 1956 (E. J.
Ford, Jr., Bishop Mus.). West N. G.: 34,
Hollandia, July-Sept. 1944 ( Darlington ) ; 6,
same locality. May 1945 (B. Malkin,
U.S.N.M.); 5, Sabron, Cyclops Mts., 930
ft. (c. 280 m), Apr. 1936 \ Cheesman ) .
Measured specimens. The £ holotype and
1 9 paratype from Dobodura.
Notes. This new species is similar to
Hyphaereon reflexus Macleay (the type
species of the genus) of Java, but timidus
differs slightb' in proportions (e.g., the
base of the prothorax is relatively narrower
than in reflexus) and the pronotum of
timidus is less extensively punctate.
The wings of this species vary (Figs. 29,
A, B), and the variation is complex, being
partly individual, partly geographic, and
partly correlated with Iwdy size. Of the
specimens from Dobodura, 5 have wings
large and folded at apex; 51, strongly re-
duced. However, both the long- and the
short-winged forms are variable in the
Dobodura series. Among the long-winged
individuals, some have wings about 10%
shorter than others and with slightly weak-
ened venation, and among the short-winged
ones, the wing vestiges vary from about %
to about V2 the length of an elytron. In
the series from Oro Bay (only a few miles
from Dobodura) the proportion of long-
and short-winged individuals is different:
10 are long-winged, 9 short-winged. Seven-
teen specimens from Nadzab and 3 from
other localities in N-E. N. G. are all fully
winged or at least have wings long and
folded at apex. But my series from Hol-
landia is again dimorphic: 7 specimens
are long-winged, 27 short-winged. All the
long-winged specimens from Dobodura, Oro
Bay, and Hollandia are large. Some short-
winged individuals are equally large, but
there is much more variation in size among
the short-winged ones. I do not remember
noting this correlation in any other Ca-
rabidae. I have not studied state of wings
in specimens not collected by myself be-
cause I do not know how they were taken,
and method of collecting may have favored
getting one wing class more than another.
It is doubtful if even the long-winged
form of this species flies. Individuals are
common at some localities where much
collecting has been done, but few have
been obtained except by myself (on the
ground), and no specimen is labeled as
taken at light. The variation and use of
wings in this species would be an interesting
subject for study in the field.
Although my field notes are scanty, I
think my series of this species were taken
among dead leaves and vegetation on the
ground near water.
68
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Hyphaereon cordons n. sp.
Description. With characters of genus;
form N ebria -\ike; black or piceous, append-
ages irregularly brown, darker than in other
species. Head 0.67 and 0.67 width pro-
thorax. ProfJiora.x cordate; width/length
1.35 and 1.43; base apex 1.23 and 1.18;
base/head 1.14 and 1.12; sides converging
and broadly sinuate before slightly obtuse
(almost right), slightly blunted posterior
angles. Elytra c. %o or more wider than pro-
thorax (elytra prothorax 1.32 and 1.32);
humeri ± subangulate; striae deep, intervals
convex. Win^is fully developed, or at least
long and folded at apex, in all specimens.
Seconckiry se.xual cJiaracters as for genus.
Measurements: length c. 7-8.5; width 2.7-
3.4 mm.
Types. Holotype £ (M.C.Z., Type No.
31,388) and 77 paratypes all from Chimbu
Vy., Bismarck Rge., N-E. N. G., 5000-7500
ft. (c. 1500-2300 m), Oct. 1944 (Dar-
lington ) .
Additional material. N-E. N. G.: 1,
Kainantu, 1650 m, Oct. 20-26, 1959 (T. C.
Maa, Bishop Mus.); 1, Wau, Morobe Dist.,
1200 m, Dec. 4-5, 1961 (Sedlacek), in
mercury vapor light trap.
Measured specimens. The <^ holotype and
1 9 paratype.
Notes. Although this species is evidently
sometimes common, and although all speci-
mens are fully winged, they are rarely
taken in light traps, which suggests that
even this winged species rarely flies.
Whether it is a real species or a local form
of timidus is not possible to say from
museum specimens. In any case it is
clearly distinguishable as indicated in the
preceding Key.
Genus ANOPLOGENIUS Chaudoir
Chaiuloir 1852, Bull. Soc. Nat. Moscow 25, 1, p.
88.
Csiki 1932, Colcop. Cat., Carabidae, liarpaliiiae 6,
p. 1236.
Schauberger 1937, Ent. Rundscbau 54, p. 272.
Basilewsky 1951, Ann. Mus. Congo Belj^c (8),
Zoo!., 9, p. 122 (see for synonymy and additional
refcronces).
Diagnosis. Relatively large Acupalpina;
anterior marginal line of pronotum entire
and deeply impressed; scutellar striae ab-
sent.
Description. None required here.
Type species. Stenoloplius alaccr Dejean,
of Africa.
Generic distribution. The warmer parts
of the Old World; in the Oriental-Aus-
tralian area, from China and Japan to
northern Australia.
Notes. The species of this genus are
among the most aquatic of Carabidae, oc-
curring as a rule in vegetation that is float-
ing in water. They are active and winged
and are common in some places, including
the Philippines, although unaccountably
rare or local in New Guinea.
Anoplogenius marginafus (Macleay)
Macleay 1888, Proc. Linnean Soc. New South
Wales (2) 3, p. 472 (HarpJancr).
?iticisus Andrewes 1926, Ann. Mat;. Nat. Hist. (9)
18, p. 279.
?poJitus Schauberger 1937, Ent. Rundschau 54, p.
273.
Description. None needed here. See
Notes below; length c. 6.5-8.0 mm.
Types. Of murginattis, from King's
Sound, Australia; probably in Macleay
Mus., Sydney (not seen). Of incisus, from
Fort de Kock, Sumatra; in Briti.sh Mus.
(seen). Of politus, from "Tigerinsel (New
Guinea)" (? = Matjan, Pulau-Pulau, which
is really not off New Guinea but south of
Celebes); in Stockholm Mus. (not seen).
Occurrence in New Guinea. The only
(supposedly) New Guinean specimens of
Anoplogenius known to me are: 1 labeled
simply "Papua," presumably collected by
Biro, now in Hungarian National Mus.;
scNcral Irom "Dorey, New (Guinea," pre-
sumabK- collected by Wallace and perhaps
actual]) from Celebes (see Part I of the
present work, pp. 330-331); and the types
ol ])olHus from "Tigerinsel," probably off
(Celebes rather (liaii New (Guinea.
Notes. The Oriental- Australian species of
Anoplogenius are taxonomicalK difficult at
b(\st, and in the case of this New (Uiinean
The Carabid Beetles of New Guinea • Darlington
69
species the difficulty is increased by lack
of adequate material and by doubt about
localities, as indicated above. The three
authors concerned published their descrip-
tions without reference to each other, but
Andrewes and Schauberger both compared
their species with A. cyanescens Hope, and
Macleay's Harplaner marginatus is appar-
ently an Anoplogenius near cyanescens (B.
P. Moore, personal communication, 1965).
I therefore tentatively conclude that incisus,
politiis, and marginatus are probably all
one species which ranges at least from
Sumatra to New Guinea and northern
Australia. This species is narrowly or
indistinctly pale-margined, with relatively
distinct (narrowly rounded) posterior pro-
thoracic angles, and with baso-lateral im-
pressions of pronotum extensively punc-
tate. I have specimens with these charac-
ters from Morotai Island in the Moluccas,
and from Townsville and Rockhampton,
Australia.
Genus EGADROMA Motschulsky
Motschulsky 1855, fitude Ent. 4, p. 43.
Csiki 1932, Coleop. Cat., Carabidac, Harpalinae 6,
p. 1239 (as subgenus of Actipalpus) (see for
additional references ) .
Jeannel 1942, Faune de France, Coleop. Carabiques,
Part 2, p. 699.
Basilewsky 1951, Ann. Miis. Congo Beige (8),
Zoo!., 9, p. 144.
Diagnosis. See Key to Genera of Harpalini
of New Guinea.
Description. None required here.
Type species. Carabus smaragduhis Fab-
ricius, below.
Generic distribution. The warmer parts
of the Old World: Africa and Madagas-
car ( 1 species reaching the Mediterranean
part of Europe); southern Asia north to
Japan, and across the Malay Archipelago
to the Philippines and Australia.
Notes. The Oriental-Australian species of
Egadroma are exceptionally difficult. They
are closely related or at least very similar
among themselves; they vary geographically
and individually; and some species are
widely distributed and very common, so
that many specimens fell into the hands of
early taxonomists who described them in-
adequately and failed to understand their
interrelationships. I do not pretend fully
to understand them now, but can make
the following comments on the species that
occur in New Guinea. Three common
species occur there, distinguishable by both
external and genitallic characters. (A fourth,
endemic species is known from a single ? . )
All three are widely distributed outside
New Guinea and at least two of them have
received different names in different places.
To fix the synonymy of these species out-
side New Guinea would be a major, time-
consuming undertaking, and is beyond my
power now. I shall therefore simply use
for each species the name that I think ap-
plies in New Guinea, with tentative notes
on synonymies.
Although I did not always distinguish
the species in the field, my notes suggest
that quinquepustulata and smaragdula oc-
cur in wet places usually by standing water,
but that robusta occurs principally in drier
habitats, especially under cover in open
grassland. All these species are winged,
and all fly to light.
Key to Species of Egadroma of New Guinea
1. Size larger (c. 6-7 mm); elytra conspicu-
ously 3- or 5-maculate, and very shining;
apex of aedeagus long (p. 70 )
quinquepustulata
- Usually smaller (less than 6 mm, except in
Cyclops); usually with reduced or no mark-
ings, and often ( not always ) less shining;
apex of aedeagus shorter (except unknown
in Cyclops) 2
2. Large (7.4 mm) (p. 70) cyclops
- Smaller ( less than 6 mm ) _., 3
3. Form relatively narrower, with relatively
wider head ( head usually more than 0.74
width prothorax, prothoracic width/length
usually less than 1.45); elytra rather strongly
iridescent; aedeagus finely notched at sides
(p. 70) smaragdula
- Broader, with relatively narrower head (head
usually less than 0.74 width prothorax, pro-
thoracic width/length usually more than
1.45); elytra less iridescent, usually with
distinct microreticulation; aedeagus not
notched at sides (p. 71) robusta
70 Bulletin Museum of Comparative Zoolofiy, Vol. 137, No. 1
Egadroma quinquepusfulafa (Wiedemann)
Spiistiilatiis- Wiedemann 1823, Zoologisches Maga-
zin (2) 1, p. 58 (Badi.stcr) .
Csiki 1932, Coleop. Cat., Caral)idae, Harpalinae
6, p. 1240 (see for additional references).
Habu 1961, in Kira and ITniesao, Nature and Life
in Southeast Asia (Kyoto) 1, p. 276, fig. 4
( <5 genitalia ) .
Description. None needed here; see pre-
ceding Key to Species and following Notes;
length c. 6-7 mm.
Type(sj. From Bengal, India; in Copen-
hagen U. Mus. (not seen).
Occurrence in New Guinea. Widely dis-
tributed at low altitudes, fairly common:
41 specimens from localities over most of
the length of New Guinea (Papua to the
Vogelkop); most at low altitudes, but 1,
Chimbu Vy., Bismarck Bge., 5000-7500 ft.
(c. 1500-2300 m), and 1, Wau, 1200 m.
Notes. This relatively large, clearly
marked, and therefore comparatively easily
recognized Efradroma ranges from SE.
Asia including Japan and Formosa across
the Malay Archipelago to North Queens-
land, Australia. In the past, the species
has often been treated as a variety of
smaraiiduki but it is unquestionably distinct
by genitallic as well as external characters.
Specimens from New Guinea vary in
elytral markings, the variation being in-
dividual, not geographic. Conspicuous post-
humeral pale spots are always present, and
a variable (sometimes Faint) subapical
sutural mark is always present too, but
antcapical spots on the 7th and 8th intervals
are variably developed and often absent.
Egadroma cyclops n. sp.
Descri])lio)i. Form as in T'igure 30, larg(\
broad; side margins of j^rothorax testaceous-
translucent, ol elytra scarcely so; append-
ages reddish testaceous, antennae darker
from 3rd segment; shining, front with iso-
diametric reticulate microsculpture, discs
of pronotum and elytra not visibly micro-
reticulale (at 5()X ) but moderately irides-
cent. Head 0.6.S width prothorax; formed
as usual in genus. Prothorax transverse;
width length 1.45; base/apex 1.22; sides
rounded, with moderate reflexed margins;
basal angles rounded; base not margined,
apical marginal line interrupted at middle;
baso-lateral impressions broad, shallow, c.
rugose-punctate, with punctation finer and
sparser at middle of base. Elytra: width
elytra/prothorax 1.33; humeri prominent
but rounded; striae deep, impunctate;
scutellar striae long; intervals finely sparsely
punctulate, 3rd with 1 puncture on inner
edge less than % from apex. Winii,s full.
Lower surface: prostemum with some short
pubescence; abdomen not pubescent at
apex. Secondary sexual cJiaractcrs: i un-
known, 9 normal. Measurements: length
7.4; width 3.3 mm.
Type. Holotype 9 (Bishop Mus.) from
Hollandia area, W. Sentani, Cyclops Mts..
West N. G., 50-100 m (Gressitt and T. C.
Maa), in light trap.
Notes. This new species resembles
smaraiidula but is larger (a large smaragdula
is less than 6 mm long), with relatively
narrower head. I feel sure it is a distinct
species although reprc-sented b\- onl>- a
single 9 specimen.
Egadroma smaragdula (Fabricius)
Fabricius 1798, SuppUineutuni Eut. Systematicac.
p. 60 (Camhits).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6.
p. 1241 (see for additional references).
Jedlicka 1935, Acta Soe. Ent. Czechoslovakia 32,
p. 1 13 (in key).
Habu 1961, in Kira and Umesao, Nature and Lite
in Southeast Asia (Kyoto) 1, p. 275, fig. 3 ( S
genitalia).
Descri))liou. None required here; see
preceding Key to Species: k-ngth ± 5 mm.
Typc(s). From "India ori«Milali"; in
CoiXMihagen V\n\. Mus. (not sec-n).
Occurrciu-e in New Guinea. Common at
low altitud(\s probably throughout New
(i^uinea: 121 specimens from Miln(> Ba\
and Port Moresb>- to the X'ogelkop; most at
low altitudes but 2 from Wau and 1 from
Rattan Ciuup at 1200 in.
Notes. This is the common, unmarked
(at most with a faint rufesc(Mit area along
The Carabid Beetles of New Guinea • Darlington 71
suture posteriorly) Egadroma of the Ori-
ental-Australian area. It apparently ex-
tends from Asia to northern Australia but,
liecause of difficulty in distinguishing it
from similar forms, I have not tried to fix
the exact limits of its range.
Egadroma robusta Sloane
Sloane 1907, Deutsche Ent. Zeitschrift for 1907, p.
469.
Andrewes 1927, Ann. Mag. Nat. Hist. (9) 19, p.
110 (as synonym of dingo Castelnau).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1242 (as synonym of vestigialis) .
Description. None required here. See
preceding Key to Species and Notes, be-
low; length ± 5 mm.
Types. From Gazelle Pen., New Britain;
should now be in Deutsches Ent. Mus.,
Berlin-Dahlem (not seen).
Occurrence in New Guinea. Very com-
mon throughout New Guinea at low alti-
tudes: 368 specimens from many localities
from Milne Bay and Normanby, Ferguson,
Rossel, and Sudest Islands, Papua, to west-
ern part of West N. G. Although this species
does not commonly occur at altitudes of
more than a iew hundred meters, I ha\e
seen 4 from Wau, 1200 m, and 1, Chimbu
Vy., 5000-7500 ft. (c. 1500-2300 m). Speci-
mens have been collected in every month.
Notes. Most individuals of this species
from New Guinea have the elytra un-
marked or with only small posthumeral
pale marks on the 6th intei^vals, rarely
extending to the 5th and 7th intervals.
Similar unmarked individuals occur in New
Britain (types of robusta) and Cape York,
Australia (collected by me in 1958).
However, a few specimens from New
Guinea, mostly from the far west including
Biak Island, have also pale subapical
sutural dashes and variable subapical pale
spots, best developed on the 7th intervals.
These specimens may be referable to
Egadroma quadrimaculata (Macleay), which
was described from Australia, but which
may extend across the Malay Archipelago
at least to Java. In other words, robusta
may be an unspotted form ( occurring prin-
cipally but not exclusively on New Guinea )
of a more widely distributed maculate
species, tentatively identified as quadrimac-
ulata Macleay. This case requires further
study.
Most individuals of robusta are easily
separable from smaragdula by proportions
(indicated in the preceding Key) and
duller surface. Apparent intermediates do,
rarely, occur. I do not know whether they
are hybrids or individual variants. These
two species probably occupy different habi-
tats: smaragdula, wet places; robusta, rel-
atively dry ones.
Genus STENOLOPHUS Stephens
Stephens 1828, iHustrations British Ent., Man-
dibulata 1, pp. 67, 165.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae
6, p. 1259 (see for synonymy and additional
references )
Jeanne] 1942, Faune de France, Coleop. Carabiques,
Part2, p. 693.
Basilewsky 1951, Ann. Mus. Congo Beige (8),
Zool., 9, pp. 118, 213 (in text).
Diagnosis. See Key to Genera of Harpalini
of New Gui7iea.
Description. None required here.
Type species. Carabus teutonus Schrank,
of Europe, etc. (see Jeannel, 1942).
Generic distribution. Eurasia and North
America, and probably the Oriental Re-
gion and Malay Archipelago to Australia;
(probably not Africa below the Sahara, al-
though closely related genera occur there).
See Notes, below.
Notes. Jeannel and Basilewsky have
divided Stenolophus, and Basilewsky sug-
gests that the genus in a strict sense may be
confined to the Holarctic Regions. How-
ever, the following t\\'o New Guinean
species seem to fit reasonably well in
Stenolophus according to characters given
by Basilewsky (1951, p. 213). Moreover,
one of these two species (gonidius) has
the first segment of the posterior tarsi
plainly carinate while the other (volucer)
has not, which suggests that this character,
which has been used in dividing St e nolo-
72 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
phus, is less important than has been
thought. I shall therefore leave both New
Guinean species in Stenolophus, where in
fact earlier authors put them.
Both the following two species are ap-
parently widely distributed in the Malay
Archipelago, and I have specimens prob-
ably representing both from North Queens-
land, Australia, but because of doubt about
identifications I prefer not to state their
distributions in detail.
Key to Species of Stenolophus of
New Guinea
1. Sides of prothorax sinuate before base, with
liasal angles nearly right and scarcely blunted
(p. 72) volucer
- Sides of prothorax c. straight, converging
but not or scarcely sinuate posteriorly, with
basal angles obtuse, narrowly rounded (p.
72 ) gonidiiis
Stenolophus volucer Andrewes
Andrewes 1930, Arkiv for Zoologi 21A, No. 29, p.
5.
Description. None required here; length
c. 5.5-6.0 mm.
Types. Five, from Sumatra; actual
"type" in Stockholm Mus. (not seen).
Occurrence in Neio Guinea. Papua: 52
specimens from various localities, including
a series from Dobodura. N-E. N. G.: 1,
Nadzab, Julv 1944 (Darlington). West
N. G.: 1, River Tor (mouth) 4 km E.
Hoi Maffen, July 19, 1959 (T. C. Maa,
Bishop Mus.).
Notes. See Notes under the genus. I
have identified this species from descrip-
tion, and am not ((uite sure of it. It is, as
Andrewes says, about the si/c and color of
ii^onidius (below) but with lateral borders
of elytra darker and with differently formed
prothorax.
Stenolophus gonidius Bates
Bates 1890, Ann. Mus. C:iv. Ceiioa 27, p. 101.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1262 (see for additional references).
Andrewes 1933, Cat. Carabidae Sumatra, p. 317.
Description. None required here- l(>ngth
c. 5.5-6.5 mm.
Types. From Burma; in Genoa Mus.
(not seen).
Occurrence in New Guinea. Papua: 31
specimens from Dobodura, Oro Bay, Port
Moresby, Fly R., and Ferguson Is. N-E.
N. G.: 1, Finschhafen, Huon Pen., 80 m,
Apr. 16, 1963 (Sedlacek), in Malaise trap.
( No specimens from ^^^est N.G. )
Notes. See Notes under the genus. Al-
though I have seen a cotype of gonidius
in the British Museum and have made com-
parisons with it, I am not quite sure of the
identity of the New Guinean specimens.
Genus ACUPALPUS Latreille
Latreille 1829, in Cuvier, Regne Animal, ed. 2,
4, p. 391.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
pp. 1238, 1242.
Jeannel 1942, Faune de France, Coleop. Carabiqucs,
Part 2, p. 712.
Basilewsky 1951, Ann. Mus. Congo Beige (8),
Zool., 9, pp. 232, 233.
Diap,nosis. See preceding Key to Genera
of UarpaJini of Neic Guinea and following
Notes. I ha\'e tentatively assigned to this
genus all the small Harpalini of New
Guinea ( 6 species ) that possess lonii,
sparse prosternal setae. All are winged.
The front tarsi of the S are not or at most
(in furvinus) slightly dilated, with squamae
thin, transparent, difficult to detect, and
perhaps absent in some cases.
Description. None required here.
Type .■species. Carahus nicridianus Lin-
naeus, of Europe.
Generic distribution. Now considered to
include all principal regions of the world.
but the generic classification of these small
Harpalini is not lully worked out. At least
3 stocks of this genus reach nortliern tropi-
cal Australia, but the\ aiiparenth' do not
extend far into southern tempcMate Aus-
tralia, where tluMr place is taken In small
species of Lccanonwrtis (see discussion un-
der Tribe Harpalini).
Notes. Several species ol this genus (>ither
range widely in the Oriental-Australian area
or b(4()ng to wide-ranging groujis of closely
interrelaled species. Their nomenclature is
The Carabid Beetles of New Guinea • Darlington 73
difficult, and synonymies outside New
Guinea remain to be determined.
Key to Species of Acu palpus of
New Guinea
1. Anterior margin of clypeus not notched or
impressed at sides ( at most slightly sinuate ) ;
base of prothorax c. squarely truncate, with
posterior angles c. rectangular 2
- Anterior margin of clypeus notched or im-
pressed at sides; posterior angles of prothorax
usually more obtuse or rounded . 3
2. Larger (c. 3.7 mm); elytral margins behind
humeri faintly serrate at 50 X ( P- 73) __ cxactus
- Smaller (c. 2.7 mm); elytral margins not
visibly serrate at 50x (p- 73) -- exactellus
3. Prothorax with baso-lateral impressions not
punctate; posterior angles (narrowly) roimded
(p. 74) ftirvinus
- Prothorax with baso-lateral impressions punc-
tate; posterior angles usually more distinct ^ 4
4 Color brown with darker head; prothorax
with base more oblique at sides and posterior
angles more obtuse (p. 74) hrunnicolor
- Color darker, more uniform; prothorax with
base less oblique at sides and basal angles
more nearly right 5
5. Smaller (3.0-3.3 mm) (p. 75) ustus
- Larger (3.5-4.2 mm) (p. 75) papiia
Acupalpus exactus n. sp.
Description. With characters of genus;
form ( Fig. 31 ) relatively slender, elytra
subparallel; piceous, clypeus etc. reddish,
margins of prothorax rather broadly and
indefinitely reddish, suture and margins of
elytra narrowly but conspicuously reddish,
appendages testaceous; shining, dorsal
microsculpture faint or absent. Head 0.84
and 0.85 width prothorax; eyes large, promi-
nent; frontal impressions deep, converging
anteriorly, ending at deep clypeal suture;
anterior margin of clypeus slightly sinuate
but not distinctly notched at sides; man-
dibles long; mentum not toothed; ligula
slender, free at apex, 2-setose; paraglossae
slightly longer than ligula, narrowly
rounded; palpi with apical segments sub-
conical. Prothorax broadly subcordate;
width/length 1.34 and 1.30; base/apex 1.10
and 1.07; sides broadly rounded anteriorly,
slightly converging and broadly sinuate to
c. right, sharply defined posterior angles;
base and apex not margmed; lateral mar-
gins moderate, not crenulate; baso-lateral
impressions large, deep, irregular but not
distinctly punctate; disc normal, impressed
median line groove-like at base. Elytra
long; width elytra/prothorax 1.47 and 1.52;
humeri prominent but rounded; margins
behind humeri visibly serrate (at 50x);
striae deep, entire; intervals convex, 3rd
with puncture on inner edge well behind
middle. Measurements: length c. 3.7;
width 1.4-1.5 mm.
Types. Holotype 9 (M.C.Z., Type No.
31,389) from Hollandia, West N. G., July-
Sept. 1944 (Darlington); and 1 9 paratype
(Bishop Mus.) from Kiunga, Fly R., Papua,
July 15-21, 1957 (W. W. Brandt); 1 para-
type, Popondetta, Papua, 25 m. May 1966
( Shanahan-Lippert, Bishop Mus.), light
trap.
Notes. This New Guinean species resem-
bles and is probably related to Acupalpus
horni Andre wes of SE. Asia but is darker,
with lateral margins of prothorax not
crenulate (faintly crenulate in horni), and
with baso-lateral impressions of pronotum
less linear. Other apparently related forms
occur in the Philippines, Moluccas ( Morotai
Is.), and North Queensland in Austraha.
Acupalpus exactellus n. sp.
Description. With characters of genus;
smaller and relatively shorter than exactus
( above ) ; piceous, margins of prothorax and
elytra not or not conspicuously paler, ap-
pendages testaceous; moderately shining,
reticulate microsculpture (slightly trans-
verse ) visible on front anteriorly but absent
on discs of prothorax and elytra. Head 0.75
and 0.76 width prothorax; eyes relatively
much smaller than in exactus, with genae
more oblique; front similarly impressed;
clypeus without distinct notches at sides
anteriorly; mouthparts as in exactus. Pro-
thorax broadly subcordate with wide base;
width length 1.40 and 1.42; base/apex 1.24
and 1,16; sides broadly arcuate, slightly
converging and broadly sinuate to c. rec-
tangular posterior angles; base and apex
not margined; lateral margins narrow an-
74 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
teriorly, l:)ioader posteriorly, not crenulate;
baso-lateral impressions broad, deep, ir-
regular, but not distinctly punctate; disc
normal, with middle line deeply impressed
basally. Elytra: \\'idth elytra prothorax
1.41 and 1.45; humeri broad but rounded;
margin behind humeri not visibly serrate
(at 50x ); striae and dorsal punctures as in
exacttis. Secondary sexual characters: S
front tarsi not dilated and apparently with-
out squamae ( see Notes, below ) . Measure-
uicuts: length c. 2.7; width 1.3 mm.
Types. Holotype $ (M.C.Z., Type No.
31,390) and 1 9 paratype from Hollandia,
West N. G., July-Sept. 1944 (Darlington).
Notes. So far as I know, the only species
with which the present one need be com-
pared is exacttis (above), with which com-
parison is made in the Key and preceding
Description.
Although the type is a $ ( dissected ) , the
anterior tarsi are slender and without the
expected squamae, so far as I can deter-
mine under highest power of my stereo-
scopic microscope.
Acupalpus furvinus n. sp.
Descri])tion. With characters of genus;
form rather elongate; reddish piceous,
sides of prothorax and suture and sidt\s of
elytra morc^ reddish, appendages irregularly
testaceous, antennae brownish except at
base; shining, but front with light iso-
diametric microsculpture, pronotum with
more transverse microreticulations at most
hiintly indicated, elytra with distinct trans-
verse meshes (at 50x ) but sHghtK' irides-
cent. Head 0.71 and 0.73 width prothorax;
eyes moderate; front impressed as usual;
clypeus with anterior margin notched or
impressed at sides; mouthparts as in
e.xactus. Prothorax subquadrate; width
length 1.31 and 1.31; base/apex 1.10 and
1.11; sides broadly rounded anteriorly,
nearly straight (or slightly rounded) and
converging posteriorly to narrowly round(>d
posterior angles; base and apex not mar-
gined; margins moderate, broader jioste-
riorl\ and rniming into irregular, rounded.
not distinctly punctate baso-lateral impres-
sions; disc as usual. Elytra: width elytra/
prothorax 1.38 and 1.39; striae entire, well
impressed; 3rd intervals 1-punctate on inner
edge behind middle. Secondary .sexual char-
acters: i front tarsi slightly dilated, 4th
segments deeply emarginate, 4 segments 2-
seriately squamulose. Measurements: length
3.5-4.7; width 1.5-1.8 mm.
Types. Holotype $ (M.C.Z., Type No.
31,391 ) and 6 parat>'pes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. Papua: 1,
Lake Daviumbu, Flv R., Sept. 21-30, 1936
(Archbold Exp., A.M.N.H.). N-E. N. G.:
1, Aitape, Aug. 1944 (Darlington).
Measured s))eci7nens. The 6 holotype and
1 9 paratype from Dobodura.
Notes. According to my notes made at the
British Museum in 1948, this species rep-
resents a widely distributed Oriental-Aus-
tralian group of Acupalpus which includes
annarncnsis Bates and jurvus Andrewes of
SE. Asia, as well as various named forms
from the Malay Archipelago. Most of them
are testaceous with dark elytral clouds, but
jurvus is more uniformly colored, as is
furvimis. However, the latter differs from
jurvus in haxing more prominent eyes and
less rounded posterior prothoracic angles.
A representati\ e of this species group oc-
curs in North Queensland, Australia.
Acupalpus brunnicolor (Sloane)
Sloane 1S98, Proc. Linnean Soc. New South Wales
23, p. 466 (Thcnarotcs).
Andrewes 1930, Cat. Indian Inseets, Part 18,
Caral)i(Iai', p. 10.
Description (for recognition only). A
brown Acupalpus characterized in preced-
ing Key to Species: head 0.82 and 0.82
widtli prothorax; prothoracic width length
1.44 and 1.42, base apex 1.13 and 1.16;
width elytra prothorax c. 1.43 and 1.38;
length c. 3.8, width 1.5-1.6 nun.
Types. From Behn Ri\er, Wcslcru Aus-
tralia, collected 1)\' Helms; t\pe retinned
to Lea, should be in Sonlh Australian Mns.
(not secMi).
The Carabid Beetles of New Guinea • Darlington 75
Occurrence in Neiv Guinea. Papua: 3,
Port Moresby, Oct. 1944 (Darlington); 1,
Orionio River, Feb. 17, 1967 ("H. C,"
Bishop Mus.), light trap.
Measured specimens. A pair ( c5 9 ) from
Port Moresby.
Notes. According to notes that I made
at the British Museum in 1947-1948, brun-
nicolor of Avistralia may be a form of a
widely distributed species that has received
the name sinuellus Bates in SE. Asia and
punctafus Jedlicka in the Philippines. How-
ever, the classification of these small har-
palines is still so doubtful that I do not
wish to declare synonymies, but shall say
only that hrunnicolor probably represents a
widely distributed species group that may
have reached Australia recently (perhaps
by way of the Lesser Sunda Islands) and
that may then have spread from Australia
to the Eucalyptus country of southern New
Guinea.
Acupalpus ustus Andrewes
Andrewes 1930, Zool. Mededelingen 13, p. 195.
Description ( for recognition only ) . With
characters of genus; form rather stout;
piceous, moderately shining. Head 0.76
and 0.77 width prothorax; eyes average.
Prothorax subcordate; width/length 1.38
and 1.37; base apex 1.12 and 1.10; sides
variably sinuate before obtuse (almost
right) posterior angles; disc normal, baso-
lateral impressions punctate, punctation ab-
sent or sparse across middle of base. Elytra:
width elytra/prothorax 1.43 and 1.44. Mea-
surements (in New Guinea): length 3.0-
3.3; width l..'3-1.4 mm.
Types. From Sumatra and Borneo; the
(holo)type, from Borneo, in Andrewes
Coll., British Mus. (seen).
Occurrence in Neiv Guinea. Papua: 29,
Dobodura, Mar.-July 1944 (Darlington).
West N. G.: 1, Hollandia, July-Sept. 1944
( Darlington ) .
Measured specimens. A pair { i 9 ) from
Dobodura.
Notes. My identification is tentative, al-
though based on comparison of specimens
with the type. As in other species of this
genus, I am not absolutely sure of synon-
ymies and make none, but only state that
this species appears to be widely distributed
in the Malay Archipelago.
Acupalpus papua n. sp.
Description. With characters of genus;
form (Fig. 32) rather broad, with sides of
elytra shghtly arcuate; piceous, apices of
elytra sometimes rufescent, appendages
testaceous, antennae browner from 3rd seg-
ments; moderately shining, reticulate micro-
sculpture distinct and c. isodiametric on
front, faint ( and more transverse ) or absent
on disc of pronotum, not visible (at 50x )
on slightly iridescent elytra. Head 0.82 and
0.80 width prothorax; eyes moderate; frontal
impressions extending onto clypeus; anterior
edge of clypeus finely notched at sides;
mouthparts c. as in exactus. Prothorax
broadly subcordate; width/length 1.43 and
1.46; base/apex 1.15 and 1.10; sides broadly
rounded anteriorly, converging and slightly,
broadly sinuate before slightly obtuse but
well defined posterior angles; margins
moderate anteriorly, broader posteriorly,
with baso-lateral areas depressed and punc-
tate, punctation not extending across middle
of base. Elytra: \\'idth elytra/prothorax
1.46 and 1.48; humeri broadly rounded;
margins behind humeri not distinctly serrate
at 50X; striae moderately impressed; 3rd
intervals 1-punctate on inner edge as usual.
Measurements: length 3.5-4.2; width 1.5-
1.8 mm.
Types. Holotype S (M.C.Z., Type No.
31,392) and 4 paratypes from Dobodura,
Papua, Mar.-July 1944 (DarHngton); and
additional paratypes as follows. Papua: 1,
Oro Bay (near Dobodura), Dec. 1943-Jan.
1944 (Darlington); 1, Fly R., Lake Davi-
umbu, Sept. 1-10, 1936 (Archbold Exp.,
A.M.N.H.). West N. G.: 2, Hollandia,
July-Sept. 1944 (Darlington).
Measured specimens. The c^ holotype and
1 ? paratype from Dobodura.
76 BuUetin Museum of Comparative Zoology. Vol. 137, No. 1
Notes. Except that it is close to tistiis
( above ) , I cannot state the relationships of
this new species. As compared with iistu-s,
papim is larger, with relatively slightly
w ider head and wider prothorax.
Tribe ANAULACINI
Csiki 1932, Coleop. Cat. Carabidae, Harpalinae 7,
p. 1287 ( see for synonymy and additional refer-
ences ) .
Jedlicka 1963, Ent. Abhandlungen 28, p. 283.
MiLsoreini Auct.
Andrewes 1930, Cat. Indian Carabidae, p. XIII.
Jeannel 1949, Coleop. Carabiques de la Region
Malgache, Part 3, p. 860.
Masorcitdc jeannel 1942, Fainie de France, Coleop.
Carabiqnes, Part 2, p. 1012.
Ma.sorcinac Basilewsky 1953, Exploration Pare Na-
tional rUpeniba, Fasc. 10, Carabidae, pp. 17,
118.
This is a small, pan-tropical tribe of
obscure ('arabidae some of which, includ-
ing all those found in New Guinea, super-
ficially resemble Nitidulidae. They may be
recognized by appearance ( Figs. 33, 34 ) ;
short, strongly arcuate, flattened mandibles;
rather long tibial spurs; and other technical
characters given by authors cited above.
y\ll the species that I know are winged,
and the widely distributed Aephnidius
adelioides flies to light, but Odontomasoreus
has not been taken at light and perhaps
do(\s not fly. The few species that I have
collected in New Guinea and Australia live
in leaf litter on the ground in rain forest.
Four genera of the tribe are known from
New Guinea.
Kkv to Genera of Anaulacini of
New Guinea
1. iiimicii dentate; (antennae short, not reacli-
ing base of prothorax; labrnni rounded;
mentnni lobed or snlidentate; chtra pale-
spotted) (p. 76) Odoiilomd.soreus
- Ilnnieri not dentate 2
2. Antennae sliort, not reaching base of pro-
tliorax (p. 77) Anaiilcicu.s
- Antennae longer, reaching or passing base
of prothorax 3
3. Mcntuiii not loolhed; size larger, c. 5 mm
(p. 77) Acp]iniditis
- Mentuni toothed: smaller, less than 3 mm
(p. 78) Caphoia
ODONTOMASOREUS n. gen.
Diagnosis. Rather small, convex Anau-
lacini; immediately distinguished from other
genera of tribe by humeri dentate.
Description. Form nitiduloid (Fig. 33),
convex; color piceous with pale elytral
marks; reticulate microsculpture isodiamet-
ric on head, slightly and irregularly trans-
verse on pronotum and elytra. Head nor-
mal, c. as in Aephnidius adelioides except
labrum rounded; antennae short, reaching
r. middle of prothorax, with median seg-
ments wider than long; mentum blimtly
toothed or obtusely prominent at middle.
Prothora.x normal; anterior angles mod-
erately advanced; base sinuate but scarcely
lobed; disc with fine, abbreviated middle
line. Elytra: humeri finely dentate; mar-
gins weakh' sinuate near apex; striae in-
dicated but scarcely impressed, scutellar
striae not visible. Secondary .sexual char-
acters: i tarsi slightly dilated, 3 segments
2-seriately squamulose; i copulatory organs
as in Figure 175.
Ty))e .species. Odontomasoreus humcralis
( below).
Generic distribution. Known (^ily from
New Guinea, thus far only from the east-
ern and central parts of the island.
Notes. I recognize only 1 species of this
new genus, with 2 subspecies.
Key to Subspecies of
Odontomasoreus fii mfrai.is
1 . Larger ( 3.4-4.0 mm ) ; humeri broadh pale
(Papua) (p. 76) humcralis ,9.,9.
- Smaller (3.1-3.5 mm); humeri dark or at
most with \ague or small pale areas (N-E.
N. G. and eastern \\'e.st N. G.) (p. 77) ^
subsp. rcdurliis
Odonfomasoreus humeralis n. sp.
Descri])lion. With cliaraclers o\ genus;
form as in Figure 33; brownish piceous,
humeri, base of elytra, and an elongate mark
on 2n(l inter\al of each elytron near apex
testaceous; mouth parts and ai)pendages
brownish or testaceous. Jlcad 0.60 and
0.63 width prothorax. ProlJiorax widest
near base, narrowed anteriorh-; width/
The Carabid Beetles of New Guinea • Darlington 77
length 1.81 and 1.81; base/apex 1.42 and
1.39; sides weakly arcuate, very narrowly
margined, each with usual 2 setae, at base
and c. -i from apex; base finely margined,
apical marginal line weak or interrupted
at middle; disc vaguely impressed each
side before base. Elytra: width elytra/
prothorax 1.10 and 1.10. Measurements:
length 3.4-4.0; width 1.7-1.8 mm.
Types. Holotype $ (M.C.Z., Type No.
31.393) and 21 paratypes all from Dobo-
dura, Papua, Mar.-July 1944 (Darlington).
Measured specimens. The i holotype and
1 9 paratype.
Notes. Most specimens of the type series
were taken in flood water in rain forest
after heavy rainfall. The insect evidently
inhabits leaf litter and perhaps loose soil
in rain forest.
Odonfomasoreus humeralis reductus
n. subsp.
Description. Similar to typical humeralis
but smaller, with the basal pale areas of
elytra reduced or absent but subapical
marks distinct. Head 0.62 and 0.62 width
prothorax. Prothorax: width/length 1.80
and 1.86; base/apex 1.45 and 1.41. Elytra:
width elytra prothorax 1.08 and 1.08. Mea-
surements: length 3.1-3.5; width 1.5-1.6
mm.
Types. Holotype $ (M.C.Z., Type No.
31.394) and 6 paratypes from Hollandia,
West N. G., July-Sept. 1944 (Darlington).
Additional paratypes from N-E. N. G.: 1,
Astrolabe Bay, 1898 ( Biro ) ; 5, Aitape, Aug.
1944 (Darlington).
Measured specimens. The $ holotype and
1 9 paratype from Hollandia.
Genus ANAULACUS Macieay
Macleay 1825, Annulosa Javanica, p. 22.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1292 (see for additional references).
Jedlicka 1963, Ent. Abhandlungen 28, p. 286.
Diagnosis. See preceding Key to genera.
Description. None required here.
Type species. Anaulacus sericeipennis
Macleay, of Java.
Generic distribution. Tropical Asia to
the Philippines and New Guinea; South
Africa.
Notes. One species (in fact, only 1 in-
dividual) of this genus has been taken in
New Guinea.
Anaulacus siamensis Chaudoir
Chaudoir 1876, Bull. Soc. Nat. Moscow 51, Part 2,
p. 25.
'hterhai Jedlicka 1934, Sbornik Ent. Odd. Nar.
Mus. Prague 1934, p. 119.
?kendengensis Louwerens 1952, Treubia 21, p. 215.
Description ( for recognition only ) . With
characters of genus; fonn (Fig. 34) of
Aephnidius but antennae relatively short;
reddish piceous without well defined mark-
ings. Head 0.66 width prothorax. Prothorax:
width/length 1.74; base/apex 1.34. Elytra:
width elytra/ prothorax 1.08. Measurements:
length c. 4.5; width c. 1.9 mm.
Type. From Siani; in Oberthiir Coll.,
Paris Mus. (not seen).
Occurrence in Neiv Guinea. West N. G.:
1, Geelvink Bay, 1878 (Raff ray & Maindron,
Paris Mus.).
Notes. This individual is identified from
description and from notes, made at the
British Museum in 1947-1948, on a speci-
men from the Andaman Islands identified
as siamensis by Andrewes. The unique
type of sterbai, from Malinao, Tayabas,
Philippine Is., is in the British Museum
too; it does not differ significantly from
the Andaman siamensis. I have a paratype
of kendengensis from Java, and it too is
very close to siamensis. All these names
probably apply to one species that ranges
from the southeastern corner of Asia to the
Philippines and New Guinea, but the
material seen is too limited to justify a
final decision.
Genus AEPHNIDIUS Macleay
Macleay 1825, Annulosa Javanica, p. 23.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1288 (see for additional references and list
of species).
Jeannel 1949, Coleop. Carabiques de la Region
Malgache, Part 3, p. 861.
Jedlicka 1963, Ent. Abhandlungen 28, p. 284.
78 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Diag,mms. See preceding Key to genera.
Description. None required here.
Type species. AepJmidiiis adelioides Mac-
leay (below).
Generic distribution. All principal trop-
ical and some adjacent warm-temperate
areas of world.
Notes. A single widely distributed species
of the genus occurs in New Guinea.
Aephnidius adelioides Macleay
Macleay 1<S25, Animlosa javaiiita, p. 23, pi. 1,
fig. 7.
Andrewes 1930, Cat. Indian Insects, Part IS,
Caral)idae, p. 11.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1288 (see for synonymy and additional refer-
ences ) .
Description. None required here; length
± 6 mm.
Type. From Java; in British Mus. (seen).
Occurrence in New Guinea. Widely dis-
tributed: 29 specimens from Papua (in-
cluding Dobodura), N-E. IS. G., and West
N. G., at low and moderate altitudes, up
to 1200 m (at Wau).
Notes. This species ranges from SE.
Asia, Japan, and Formosa to northern
Australia, east to the Philippines, New
Britain, and New Ireland. Seven speci-
mens from Wau and 1 from near Hollandia
are labeled as taken in light traps.
Genus CAPHORA Schmidt-Goebel
Schmidt-CJoebel 1846, l''aiinula (Coleop. Hiiiiuiniac,
p. 91.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1293 (see lor additional relerences).
Jedlicka 1963, Knt, Abliandlnni^'en 28, p. 288.
Diapiosis. Very small Anaulacini ( under
3 mm); characterized in the preceding Key
to (kniera of the tribe.
Description. None refjuired lu^e.
Type species. Capltora liuniilis Scliuiidl-
Goebel ( below ) .
Generic distrihntion. SE. Asia, Sumatra,
Java, the Philippin<>s, New (>iiinea, and
Cajie York. Australia (see following
species ).
Caphora humilis Schmidt-Goebel
Schmidt-Goebel 1846, Fannula Coleop. Birnianiae,
p. 91, pi. 3, fig. 8a-b.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1293 (see for additional references).
Jedlicka 1963, Ent. Abhandlnngen 28, p. 289.
Description. None required here. The
very small size distinguishes this species
from all other members of the tribe found
in the region in question. Length c. 2.5 mm.
Type. From Burma; should be in Prague
Mus. (not seen).
Occurrence in New Guinea. Papua: 1,
Brown River, May 24, 1956 (E. J. Ford, Jr.,
Bishop Mus.), in light trap.
Notes. This species is recorded from
India and Burma to Sumatra and Java
and occurs also on the tip of Cape York,
Australia (collected by me in 1958). My
Cape York specimens were found in the
Lockerbie rain forest, in leaf litter mixed
with bird droppings under a large tree in
which colonial birds had nested. Tlie
beetles were in company with Peri<i.ona
niiiriceps, which is often carried by com-
merce, and this suggests that CapJiora too
may sometimes be carried by man.
Tribe CYCLOSOMINI
This is another small tribe, repr(\sented
in all the warmer rt>gions of the world.
The name to use for it is doubtful but not
worth detailed discussion here. The mem-
bers of the tribe, although apparently re-
lated to Anaulacini, are superficialh' Lel)ia-
like but diffcT from Lebia iu having very
long tibial spurs. The only genus of the
tribe that occurs iu New Guinea is
Sarotlirocrepis.
Genus SAROTHROCREPIS Chaudoir
Chandoir 1876, linll. Soc. Nat. Mo.scow 51, Tart 2.
p. 76.
(>siki 1932, Coleop. ('at., ("arabidae, Harpalinae 7,
p. 1302 (see lor s\n()n\Mn\', additional refer-
ences, and list of species ) .
Jedlicka 1963, Knt. Abliandlunuen 2S. p. 290.
Diaii,)}osis. Sec undc^r tribe.
Dcscri))ti()n. None reciuired lieic.
The Carabid Beetles of New Guinea • Darlington
79
Type species. Carahus corticalis Fab-
riciiis, of Australia.
Generic distribution. Represented in
Australia by numerous, diverse species; 1
species group extends to New Guinea,
some Lesser Sunda Islands, Celebes, and
the Philippines.
Notes. Although most other genera of
this tribe are (I think) ground-living,
Sorotlirocrepis is arboreal. Many Australian
species live on the trunks of Eucalyptus
trees, but a few live in foliage, as does the
single New Guinean species. The tarsal
claws tend to vary with habitat. In the
foliage-living New Guinean species and also
its immediate Australian relatives, each
claw has several long teeth. In some Aus-
tralian tree-trunk-living forms, the claw
teeth are shorter or irregular.
Sarothrocrepis papua n. sp.
Description. Form as in Figure 35; ir-
regular bro\\'nish yello^^', elytra with vari-
able post-median dark brown mark usually
irregularly triangular or M -shaped; reticu-
late microsculpture isodiametric on front,
scarcely or slightly transverse on pronotum,
more transverse on elytra. Head 0.67 and
0.67 width prothorax. Frothorax: width
length 1.45 and 1.50; h-Ase head 1.41 and
1.41 (apex of prothorax too rounded for
exact measurement ) ; margins moderate an-
teriorly, much wider posteriorly ( as usual in
genus), each with seta at basal angle and
another c. ''.•! from apex; base finely mar-
gined, apical marginal line interrupted at
middle; median line lightly impressed, sub-
apical transverse impression weak, subbasal
transverse impression and posterior-lateral
impressions slight. Elytra: width elytra
prothorax 1.52 and 1.48; striae deeply im-
pressed, not punctate; a seta-bearing punc-
ture at base each 2nd interval, usually an
inconspicuous puncture on inner edge each
3rd interval near apex, and sometimes an
apparent minute puncture on outer edge
3rd interval c. V-i from base. Inner u'inii,s
full. Lower swface and legs: no note-
worthy characters except tarsal claws each
with 4 long teeth, the inner one smaller and
sometimes difficult to see. Secondary sexual
characters: i front tarsi slightly dilated, 3
segments 2-seriately squamulose (4th seg-
ments with soles of non-sexual adhesive
hairs in both sexes); S (not 9 ) apical
ventral segment deeply acutely notched at
middle; 1 seta each side apex last ventral
segment in both sexes. Measurements:
length 4.6-6.4 (usually 5.0-5.5); width 2.3-
3.0 (usually c. 2.7) mm.
Types. Holotype c^ (M.C.Z., Type No.
31,395) and 120 paratypes all from Dobo-
dura, Papua, Mar.-July 1944 (Darlington).
Additional material. Thirteen specimens
from other localities in Papua, N-E. N. G.,
and West N. G.; and 2 specimens from
Cape Gloucester, New Britain, Jan.-Feb.
1944 (Darlington).
Measured .specimens. The holotype and
1 9 paratype.
Notes. Sarothrocrepis papua resembles S.
fasciota Macleay of North Queensland, Aus-
tralia, but is larger, with prothorax narrower
and elytral markings slightly different.
Three similar species occur in the Malay
Archipelago. S. m-miorum Jordan, from
Tenimber (and in Andrewes Coll. also
from Sumbawa, Sumba, and Andonare Is. ) ,
has prothoracic margins narrower than
papua and the dark M-mark usually better
defined (but the mark is variable in papua).
S. javanica Van Emden has prothoracic
margins narrower and basal impressions of
pronotum more linear. And S. andrewesi
Jedlicka, of the Philippines, has the elytral
marks different (3 dark stripes on yellow
background) and basal impressions of
pronotum better defined, sublinear. These
names may all be based on forms of one
widely distributed variable species, but I
prefer to treat them as separate species
for the time being.
S. papua was very common in under-story
foliage of rain forest at Dobodura, espe-
cially in clumps of dead leaves still attached
to low branches. It is probably mainly
diurnal, although I have seen 2 specimens
taken in light traps.
80
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Tribe LEBIINI
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1305 ff. (see for synonymy and additional
references ) .
Jedlicka 1963, Ent. Abhandlungen (Dresden) 28,
p. 295.
Habu 1967, Fauna Japonica, Carabidae, Trun-
catipennes Croup, p. 57.
Lehiidae Jeannel 1942, Faune de France, Coleop.
Carabiques, Part 2, p. 1017.
Teannel 1949, Coleop. Caraliiques de la Region
Malgache, Part 3, p. 876.
Lehiinae Basilewsky 1953, Exploration Pare Na-
tional rUpemba, Fasc. 10, Carabidae, p. 184.
Lebiini are medium sized and small
Carabidae usually recognizable by general
appearance. Tlie elytra are usually (but
not always) obliquely truncate or sinuate-
truncate at apex (and often spined too),
and the insects have additional technical
characters, including genitalic ones, given
by authors cited above.
The tribe is a very large one. It is best
represented in the tropics and includes a
large proportion of arboreal forms especially
in the tropics.
Arboreal Lebiini divide into two ecologic
groups, one living on tree trunks and the
other in foliage. The tree-trunk dwellers
are numerous in rain forest and some occur
on fallen trees and logs as well as the trunks
and larger branches of living trees. Some
are nocturnal, some diurnal. Some of them
can be trapped under sacking tied around
tree trunks or laid over logs. Lebiini living
on tree trunks and logs in New Guinea
include Stcnotchis, Mi.scclus, Mimtthodcs,
C(it(isco))iis\ Pcricahis, Coptodcra (some),
Moclifhcrus, and Strickhindia. (Other lebi-
ine genera, especially PJidopldocus, A'^ono-
cliila, and Australian Demetrida, live on the
trunks of Eucalijpius trees in more open
woodland in Australia. ) The foliage dwel-
lers in New Guinean rain forest include
Aristolehia, Lehia, Dolichoctis, Cclacncphcs,
and especially New Guinean Demetrida.
(The ecology of Demetrida is discussed in
more detail in Note.s' imder the genus. )
Different foliage-inhabiting Lebiini in New
(Uiinea probabK inhabit dilfcrent sj^jccial
niches. Some species are commonly taken
by sweeping under-story plants in rain
forest but other species are not, and these
may live at higher levels in the trees. They
may be difficult to collect except when trees
are felled, unless they fly to light. Be-
sides the arboreal forms, a few Lebiini in
New Guinea ( and relatively more in colder
climates) live on the ground, especially in
leaf litter in rain forest. In New Guinea,
these include some Coptodcra, probably
Sijntomus- and Microlestes and Apri.stus, and
certainly Anomotarus and Nototonis. The\
can be collected in small numbers, labori-
ously, by sifting or in Berlese traps, or in
larger numbers and more easily by washing
out debris and loose earth from the forest
floor or by sorting and washing flood debris
from rain forest.
Most Lebiini are winged and many of
them fly actively to escape danger or to
disperse (see Notes under Catascojm.s),
and some, presumably mostly nocturnal
species, fly to light at night. The only
known New Guinean lebiine with atrophied
wings is Nototariis papua.
Although Lebiini are most numerous and
diverse in the tropics, the tribe as a whole
is virtualK' cosmopolitan. A few genera,
including Lehia and Coptodera, are very
widely distributed too, but most other
genera have restricted distributions.
Thirty genera of Lebiini are known in
New Guinea and at least 3 additional
genera probably occur. Many of the genera
are shared with, better represented in. and
probably derived from the Oriental Region.
These include Lehia, Catascopus, Coptodera.
Dolichoetis, and more than a do/en smaller
genera. Genera shared maiuK with and per-
haps derived from Australia are fewer but
include Aiionochda. TriiionotJiops, Phloeo-
earahii.s, Aiioniofarus, Xototarus. and espe-
cially Denielridu. The genera Miniithodes.
Striekkindia, and Miseelus now center on
New Guinea and may have originated there.
The onl\- l(>biine genus actualK confined
to New (Guinea is monotxpic Minitj)hJoeus.
The uenus Demetrida seems to hv in the
The Carabid Beetles of New Guinea • Darluii'ton
81
very midst of an explosive evolutionary
radiation, which is discussed under the
genus. I know no other case quite like it
among Carabidae.
The following key to the genera of
Lebiini of New Guinea is practical, not
phylogenetic. Genera that occur together
in the key are not necessarily closely re-
lated, and the key is designed for onJy the
New Guinean species of some genera. I
haN'e used form of the whole insect as a key
character of some genera. The form is
characteristic in some cases, and the form
of the whole is surely no less important
than the form of a part.
After the key, the genera are treated in
the order of the Coleopterorum Ccitalogus
(Csiki 1932), not in the order in which
they are keyed out.
An enormous supposed lebiine, Holopon-
erus godeffroiji (Fairmaire), has been de-
scribed from New Britain. I do not know
this insect, but I think it is probably not
a member of the Lebiini but of the tribe
Helluodini, under which I shall discuss it in
more detail.
Key to Genera of Lebiini of New Guinea
1. Fourth segments of hind tarsi deeply eniar-
ginate, with lobes \-2 or more total length
of segment 2
- Fourth segments of hind tarsi more shal-
lowly emarginate or subtnincate 10
2. Form usually broad Lebia-\ike ( Figs. .37-
41 ); base of prothorax ± lobed; upper sur-
face not pubescent; tarsi not puljescent
above; S middle tibiae excised on inner
edge near apex 3
- Form not LebiaAike, usually more slender;
l)ase of prothorax often ( not always ) with-
out lobe; tarsi often (not always) pubes-
cent above; S middle tibiae usually not
excised ( but tuberculate-serrate in many
Demetrida ) „ 4
3. Outer-apical angles of elytra sharply
formed and prothorax ± hemispheric; S
middle tibiae with 2 excisions on inner
edge near apex ( p. 83 ) Ari.stolehia
- Outer-apical angles of elytra either rounded
or sharply formed, but if latter, prothorax
not hemispheric (in New Guinean species);
$ middle tibiae with 1 excision on inner
edge near apex (p. 85) Lebia
4. Small (4-4.5 nun); protliorax trapezoidal,
widest at base (Fig. 87) (p. 134)
Peliocypas
- Larger; prothorax usually not as de-
scribed 5
5. Upper surface coarsely rugose and pubes-
cent; prothorax strongly lobed at base;
(form as in Fig. 42; length c. 8 mm)
( p. 89 ) Lachnodenna
- Upper surface not coarsely rugose; pubes-
cent or not, ])ut if pubescent, prothorax
not lobed at base 6
6. Fifth elytral intervals with coarse seta-
bearing puncture near base; prothorax with
( very short ) basal lobe ( form as in Fig.
95; length c. 8-9 mm) (p. 139) ____ Anclmta
- Fifth elytral intervals without coarse punc-
ture near base; prothorax usually (not
always) without basal lobe 7
7. Form (Figs. 97-109) usually slender; apex
of elytra sinuate-truncate or angulate or
spined but not broadly and strongly
rounded; tarsi pilose above (p. 140)
Demetrida
- Form usually less slender; apex of elytra
broadly and strongly rounded or weakly
sinuate-truncate with outer angles broadly
rounded; tarsi above pilose or not 8
8. Prothorax with (slight) basal lobe; tarsi
not pilose above (p. 184) Trigonothops
- Prothorax without basal lobe (but base
slightly oblique near angles); tarsi usually
pilose above (pilosity slight in some
Parena) 9
9. Upper surface not pubescent (p. 138) .. Paremi
- Upper surface pubescent ( p. 140 )
Endtjnomena
10. Form (Fig. 37) characteristic, broadly
oval with outer elytral angles sharply
formed; £ middle tibiae with 2 excisions
on inner edge near apex; ( length c. 9.5—
11.0 mm — smaller species of same genus
key out in couplet 3) (p. 83) ..^^ Aristolebia
- Form not as above; S middle tibiae usu-
ally not 2-excised _-_ 11
11. Form (Fig. 36) characteristic; small (c.
4 mm or less); upper surface pubescent;
prothorax with extra anterior-lateral setae;
brown with single broad transverse dark
band across elytra (p. 82) ._- (SomotiicJtus)
- Not as above in one or more ways 12
12. Very small (less than 4 mm); color black
or (rarely) transversely fasciate 32
- Larger or, if length less than 4 mm, with
color pattern of longitudinal lines 13
13. Form (Fig. 88) characteristic, slender,
with rounded elytral apices; mentum with-
out tooth and claws not toothed; (black;
length c. 7 mm) (p. 135) Celaenephes
- Not as above 14
82
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
14. Claws simple, not toothed; size usually
large; color often metallic _ - 15
- Claws each with several teeth; size often
(not always) smaller; color rarely metal-
lic 17
15. Form (Fig. 45) characteristic, slender,
subcylindric, with long genae and small
eyes, and with rounded-truncate elytral
apices; often (not always) only 1 seta
over eye; (length 9.5-14.5 mm) (p. 91) _.
Miscelus
- Form not as above; 2 setae over each
eye 16
16. Labrum long, notched at apex; size larger
( 8-22 mm ) ; color metallic or rarely brown,
without geometric marks (p. 101)
Catascopus
- Labrum shorter, not notched; smaller (7-
8 mm in New Guinean species); elytra
with geometric marks (p. 110) „.. Pericohis
17. Mentum not toothed 18
- Mentum toothed 19
18. Third elytral intervals with 2-4 dorsal
punctures, lint if 2, not as described be-
low; i middle tibiae usually excised on
inner edge before apex (p. 110) .. Coptodera
- Third elytral intervals with 2 minute non-
seta-bearing punctures behind middle, or
without recognizajjle dorsal punctures; $
middle tibiae not excised (p. 124)
_ Dolichoctis
19. Elytra spined 20
- Elytra not spined 21
20. Prothorax without extra lateral setae; form
(Fig. 44) not .strikingly broad (p. 90) _
Stciiotcltis
- Prothorax with many extra lateral setae;
form (Fig. 86) strikingly broad (p. 132) ..
Stricklatulia
21. i""orm ( F'ig. 43) characteristic, broad, with
broadly rounded elytral apices; (dull
black; lengtli c. lO-ll mm) (p. 90) _..
Siniiriis
- Not as above 22
22. Elytral apices very strongly sinuate-
truncate; (slender; color green, blue, or
coppery; length in New Guinea c. 8-9
mm ) ( p. 94 ) Holcodcrus
- Not as above __ __ 23
23. Form (Figs. 47-58) characteristic, very
broad, with wide head but relatively small
eyes, prothorax usually c. 2x wide as long,
elytra short-cjuadrate; (pubescence and
color diver.se; length c. 4-6.5 mm ) ( p.
95) Miuutlwdes
- Form not as above (if c. similar bul labrum
notched, .see Minupldoeiis, below) 21
24. Labrum notched at apex; pronotiun with
numerous lateral setae; ( shining black;
length 7.5-8.0 mm) (p. 117) ._.. Minuphloeti.s
- Labrum not notched; pronotum with 2 lat-
eral setae each side 25
25. Third elytral intervals with 3 or 4 dorsal
punctures or (in some Agonochila) these
pimctures lost amid other coarse puncta-
tion and short pubescence ^._ 26
- Third elytral intervals with 2 or rarely 1
dorsal punctures 27
26. Surface conspicuously short-pubescent and
(at least on elytra) roughened (p. 118) __
Agonochila
- Surface not distinctly pubescent, not
roughened; (3rd elytral intervals with 3
pimctures) (p. 122) Oxijodontus
27. Labial palpi slender 28
- Labial palpi with apical segments ±
widened, usually subtriangular 29
28. Third elytral inter\als 2-punctate; pro-
notum setulose (p. 122) Mochthcrtis
- Third elytral intervals 1-punctate; pro-
notmn not setidose ( p. 123 ) ^- (Mochtheroides)
29. Antennae and tarsi relatively short and
thick; i middle tiliiae arcuate and with
shallow excision at middle of length; (color
brown; length c. 8 mm) {p. 138)
( Ploch ion us )
- Antennae and tarsi more slender; S middle
tibiae not as described 30
30. Eyes abruptly prominent, genae short and
forming c. right angles with neck ( p.
183 ) Pldococarahus
- Eyes less prominent, genae longer and
forming obtuse angles with neck 31
31. Side pieces of metasternum long; inner
wings full; color pattern usualh' present
(p. 186) -- Anomotarits
- Side pieces of metasternum scarcely longer
tliau wide; inner wings vestigial; color r.
uniform brownish black (p. LSo) Xototcini.s
32. Claws simple, not tootlicd; ( incntuin with
entire tooth) (p. 137) A;)n.s7(/.s
- Claws each with se\ eral [ sometimes weak )
teeth .-.- 33
33. Mentum with (emarginate) tooth {p.
135) ___ _ _ Syntoimi.s
- Nhntuin not toothed (p. 136) .... Microlc'stcs
(Genus SOMOTRICHU5 Seidlitz)
Seidlitz 1887, Fauna lialtica, 2nd cil,, Cattungcn.
p. 7.
Nhiteu 1963, Ann. Mus. C.W. Genoa 74, pp. 131 ff.
(See also references under following species)
Didi^iiosis. Form as in I'imire 36. .small.
The Carabid Beetles of New Guinea • Darlington
83
subparallel, with eyes rather small and
\\idely separated (in this tribe); upper
surface pubescent; pronotum with several
strong setae each side; wings full; 4th hind-
tarsal segment weakly emarginate.
Description. None required here.
Type species. Carahus elevatiis Fabricius
(below).
Generic distribution. One species has
been dispersed over the wanner parts of
the world by man. A second species is
known only from Madagascar.
(Somotrichus elevatus (Fabricius))
Fabricius 1787, Mantissa Insectorum 1, p. 198
( Carahus).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1308 (see for synonymy and additional refer-
ences ) .
Jeannel 1942, Faune de France 40, Coleop.
Carabiques, Part 2, p. 1032.
Description ( for recognition only ) . With
characters of genus; brown with broad,
regular, darker brown fascia across middle
of elytra; length c. 3.5-4.0 mm.
Types. From tropical America; now in
Hunter Coll. (Glasgow) and Fabricius
Coll. (Kiel) (not seen).
Occurrence in New Guinea. Not recorded
but may occur.
Notes. Somotrichus elevatus is sup-
posedly native in tropical Africa but has
been carried over much of the world by
commerce. It is often found in seaport
cities. In the Malay Archipelago it has
been collected on Java, Celebes, and
Batjan ("Batchian") in the Moluccas, and
I have a specimen before me from Peleliu
in the Palau Is. Its occurrence in New
Guinea is therefore likely. It has not yet
been found in Australia.
Genus ARISTOLEBIA Bates
Bates 1892, Ann. Mus. Civ. Genoa 32, p. 428.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1308 (see for additional references, synonymy,
and list of species ) .
Jedlicka 1963, Ent. Abhandlnngen 28, p. 311.
Diagnosis. Similar to large Lehio with
prothorax c. hemispheric and outer elytral
angles sharply defined; wings full; 4th seg-
ments middle and hind tarsi emarginate or
lobed (see following Notes); claws with
9-11 long teeth in larger species but only
5-7 teeth in smaller species; c^ middle tibiae
each with 2 (not 1 as in Lebia) excisions
close together on inner edge near apex; most
other characters including those of mouth-
parts as in Lebia.
Description. None required here.
Type species. Aristolebia quadridentata
Bates, of Bunua.
Generic distribution. Southern India
(specimens in M.C.Z.), southern China,
Burma, etc., to the Philippines, New
Guinea, and the tip of Cape York, Aus-
tralia (see under A. wau, below).
Notes. The smaller species described be-
low are in some ways transitional between
Aristolebia and Lebia, but Aristolebia seems
to me to be a natural group worth distin-
guishing from Lebia, which is an enormous,
unwieldy genus. The tarsal lobes in
Aristolebia vary, but the variation shows
continuity. In large Asiatic species of the
genus the 4th segments of the hind and
middle tarsi are relatively weakly emar-
ginate. In the large New Guinean species
{papua) the lobes of the 4th segments are
rather short on the hind but longer on the
middle tarsi. And in the smaller New
Guinean species the lobes of the 4th seg-
ments are long, more than Vl' the segments'
length even on the hind tarsi, and are rela-
tively longer in capitis than in icau.
In New Guinea, Aristolebia occurs chiefly
at mid-altitudes. It is probably arboreal
(in rain forest) and probably diurnal, al-
though a few individuals have been taken
in light traps at Wau.
Key to Species of Aristolebia of New Guinea
1. Larger, 9.5-11.0 mm (p. 84) papua
- Smaller, 5.5-6.5 mm 2
2. Entirely yellow or brownish yellow; sutural
angles distinct and usually subdenticulate (p.
84 ) tvau
- Elytra dark with broad stripes or spots pale;
sutural angles (narrowly) rounded (p. 85) ..
capitis
84 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Arisfolebia papua n. sp.
Description. With characters of genus;
form as in Figure 37; black, sides of pro-
notum broadly and of elytra narrowly pale,
elytra with variable pale marks or some-
times wholly dark, lower surface, mouth-
parts, and appendages reddish brown;
rather shining, microsculpture as described
below. Head 0.76 and 0.73 width prothorax;
front irregularly slightly impressed and
rugulose anteriorly, rather sparsely punc-
tulate, with c. isodiametric microreticulation
especially posteriorly. Prothorox: width/
length 1.67 and 1.60; base/apex c. 1.95
and 1.98 (figures approximate because an-
terior angles not defined ) ; margins narrow
anteriorly, broad posteriorly, each with seta-
bearing puncture at basal angle and before
middle; base and apex with entire impressed
marginal lines; disc slightly transversely
rugulose, sparsely punctulate, in part lightly
microreticulate. Elytra ample; width elytra/
prothorax 1.70 and 1.67; outer-apical and
sutural angles acute and denticulate; striae
entire, impressed, faintly or not punctulate;
intervals with slightly transverse microretic-
ulation and sparse fine punctulation, 3rd
with 2 dorsal punctures on outer edge c. %
from base and less than Vi from apex
(slightly variable in position). Leii,.s: 4th
segments middle and hind tarsi as in Figure
166; claws broadly triangular, each with c.
10 long teeth. Secondary sexual characters:
6 front tarsi scarcely dilated but with nar-
row, irregularly 2-seriate squamules; i mid-
dle tibiae 2-excised; i with 2, 9 c. 4 setae
each side before apex last ventral segment.
Measurements: length 9.5-11.0; width 4.5-
5.3 mm.
Types. Holotype i (Bishop Mus.) and
17 paratypes (some in M.C.Z., Type No.
31,396) from Wau, Morobe Dist., N-E.
N. G., 1100 to 1300 m, dates in Jan.. Feb.,
Apr., May, Aug., and Sept., 1961-1963
(Sedlaceks) (holotype, 1200-1300 m. May
7, 1963); and additional paratypes as fol-
lows. N-E. N. G.: 1, Swart Vy., Karubaka,
1500 m, Nov. 11, 195S (Gressitt). West
N. G.: 1, -'Humbolt Bay" (N. A. Doherty,
British Mus.).
Additional material. Papua: 1, W . Dis-
trict, Oriomo Govt. Station, Oct. 26-28,
1960 (Gressitt).
Measured specimens. The i holotype and
1 9 paratype from Wau.
Notes. This may prove to be a geographic
form oi Arisfolebia davaonis (Heller) of the
Philippines, but the color is different
( davaonis has the prothorax rusty red, not
black) and other details are probably dif-
ferent, although I cannot be sure about
them from Heller's description of his single
specimen. A form of davaonis, or a related
species, has been found also on Salajar Is.
off Celebes (specimens received from
Louwerens ) .
The single individual from Papua is the
only one in the New Guinean series with
whollv dark elvtra, and it differs slightly
from the type series in other ways. It may
prove to be a distinguishable geographic
form. Other variation in elytral pattern is
individual in the series from Wau.
Arisfolebia wau n. sp.
Description. With characters of genus;
form c. as in preceding species {))apua):
usually entirely reddish yellow, rarely with
faint dusky areas especially at base of
elytra; upper surface with light, irregular,
c. isodiametric or slightK- transverse micro-
reticulation. Head 0.80 and 0.77 width
prothorax. Prothorax: width length 1.56
and 1.67 (difference due parth to slight
abnormal extension of basal lobe in the first
individual); lateral margins wide, flattened
or weakly reflexed especially posteriorly,
each with usual 2 setae; base and apex
with (>ntire marginal lines, but apical line
weak at middle; disc irregularh' ± trans-
versely rugulose. Elytra ample, convex;
width elytra i)rothorax 1.72 and 1.76; outer-
apical angles obtuse but well defined and
sometimes subdenticulate, sutural angles
slightK dehiscent, ± subdenticulate; striae
entire, impressed, not distinctly punctulate;
The Carabid Beetles of New Guinea • Darlington
85
intervals convex, 3rd with 2 inconspicuous
punctures on outer edge before middle and
c. Vi from apex. Legs: 4th segments hind
tarsi deeply emarginate but lobes shorter
than usual in Lebia, 4th segments of middle
tarsi with longer lobes; claws each with c.
7 long teeth. Secondary sexual characters:
i front tarsi with slender squamae probably
in 2 series but often disarranged; S middle
tibiae with 2 excisions close together on
inner edge near apex; c^ with apparently 2,
9 3 setae each side near apex last ventral
segment. Measurements: length 5.5-6.5;
width 2.7-3.2 mm.
Types. Holotype i (Bishop Mus.) and
22 paratvpes (some in M.C.Z., Type No.
31,397) all from Wau, Morobe Dist., N-E.
N. G., 1100-1500 m, dates in Jan., Feb.,
Mar., Apr., May, June, Sept., Nov., Dec,
1961-1963 ( Sedlaceks ) .
Measured specimens. The S holotype
and 1 9 paratype.
Notes. I took a single 9 of this species
at Lockerbie, near the tip of Cape York,
in January 1958, thus extending the known
range of Aristolchia to Australia.
Aristolebia capitis n. sp.
Description. With characters of genus;
fonn and characters as in wau (above)
except slightly wider; elytra dark with
either large humeral and smaller subapical
marks reddish yellow or very broad reddish
yellow stripes running from humeri to apex.
Head 0.72 and 0.72 width prothorax. Pro-
thorax: width/length 1.64 and 1.59. Elytra:
width elytra/pro thorax 1.60 and 1.69; outer-
apical angles sharply defined but sutural
angles narrowly rounded. Legs with 4th
segments middle and hind tarsi strongly
lobed (Fig. 167); claws each with 5 long
teeth and sometimes a 6th (inner) tooth
that is difficult to see. Measurements:
length c. 6.0-6.5; width 3.1-3.3 mm.
Types. Holotype S (A.M.N.H.) and 1
9 paratype (M.C.Z., Type No. 31,398)
both from Mar Village, west Vogelkop,
West N. G., Nov.-Dec. 1944 (V. S.
Mallory ) .
Notes. The rounding of the sutural angles
and the small number of claw-teeth of
capitis are Le1)ia-\ike, but the form is that
of an Aristolchia ( some Lehia approach
this form too ) and the $ middle tibiae are
decisively 2-excised.
Genus LEBIA Latreille
Latreille 1802, Hist. Nat. Crustaceorum et In-
sectorum 3, p. 85.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1310 (see for additional references, synonymy,
subgenera, and list of species).
Jeannel 1942, Faune de France, Coleop. Carabiques,
Part 2, p. 1028.
Jedlicka 1963, Ent. Abhandlungen 28, p. 314.
Diagnosis. See Key to Genera of Lebiini
of New Guinea.
Description (characters of New Guinean
species only). Form broad but variable
(Figs. 38-41); coloration variable, upper
surface not pubescent. Head: eyes promi-
nent, genae short; 2 setae over each eye;
clypeus transverse, truncate or broadly
slightly emarginate, with 1 seta-bearing
puncture each side; labrum ± transverse,
sometimes slightly rounded anteriorly, 6-
setose; mentum strongly toothed; ligula
rather broad, 2-setose; paraglossae attached
to ligula, broad, setose. Prothorax ± lobed
at base, anterior angles broadly rounded
(so base/apex ratio not determinable);
lateral margins broad posteriorly, ± reflexed,
each with 2 setae, at or near basal angle and
before middle; disc with usual impressed
middle line, weak (or obsolete) anterior
and deeper posterior transverse impressions,
and weak transverse strigulation. Elytra
wide but variable in form; striae entire; 3rd
interval 2-punctate on outer edge. Inner
wings full. Lower surface with some short,
often inconspicuous pubescence. Legs: 4th
segments middle and hind tarsi very deeply
emarginate, with long lobes; 5th segments
with accessory setae; claws with 4-6 long
teeth. Secondary sexual characters: S an-
terior tarsi not or scarcely dilated, with
slender squamae in 2 series, often disar-
ranged; S middle tibiae with 1 small deep
86
BiiUetin Museum of Comparative Zoology, Vol. 137, No. 1
excision on inner edge just before apex; $
with apparently 1 or 2 (rarely 3), 9 2 or
more seta-bearing punctures each side near
apex last ventral segment (but these punc-
tures often difficult to identify amid other
punctures and pubescence ) .
Type species. Carahus haemorrhoidaJis
Fabricius {— Lehia maiiiinato (Fourcroy)),
of Europe.
Generic distribtition. Nearly world-wide
except absent in some cold regions and on
some remote islands. Species are numerous
in most tropical regions but are relatively
few (7) in New Guinea and still fewer in
Australia. Tliis suggests that the genus
has entered the Australian Region recently,
from the direction of tropical Asia.
Notes. The New Guinean representatives
of this huge, widely distributed genus are
probably all arboreal. Some or all of them
live in the lower foliage of rain forest. They
are probably diurnal, being rarely taken
in light traps.
Key to the Species of Lebia of New Guinea
1. Outer-apical angles of elytra well defined ._^_ 2
- Outer-apical angles of elytra rounded 3
2. Color piceous; form of Endynoiucna (Fig.
3cS) (p. 86) ciulynoiuena
- Color yellow; form more of typical Lcbia
(Fig. 39) (p. 86) externa
3. Elytra with conspicuous black "anchor"
mark on testaceous background, or dark with
anterior lunules and apex testaceous (p.
87 ) ____ karenia
- Elytra differently marked or not marked 4
4. Elytra dark with large conunon cordate
area testaceous (p. 87) cordifcr
- Not tlius marked . 5
5. Brown, elytra sometimes vaguely darker or
with vague discal cloud hut not sharply
bicolored, and head and pronotum not or
only lightly microrelieulate (p. 88) i>(i])iicUii
- Eillier hicolored or with head and pronotum
heavily microreticulate 6
6. Not sharply hicolored, brown, elytra often
with disc darker; head and pronotum heavily
microreticulate (p. 88) harda
- Bicolored, head and prothorax red-testace-
ous, elytra entirely black or piceous; liead
and pronotimi not or lightly microreticulate
(p. 89) _ iii.siihinni)
Lebio endynomena n. sp.
Description. With characters of genus;
form (Fig. 38) more of Endynomena than
of typical Lebia; piceous, reflexed margins
of prothorax and ( narrowly ) of elytra trans-
lucent testaceous, appendages reddish tes-
taceous; shining, reticulate microsculpture
absent or faint on front and pronotal disc,
distinct and strongly transverse on elytra.
Head 0.82 width prothorax; front weakly
impressed at middle and on each side an-
teriorly, irregularly rather sparsely punc-
tate. Prothorax subcordate; width/length
1.69; base apex not determinable; base and
apex margined. Elytra c. % wider than pro-
thorax, narrowed anteriorly; width elytra
prothorax 1.72; apices slightly obliquely
sinuate-truncate with outer angles well de-
fined and almost subdenticulate and sutural
angles irregularly narrowly rounded; striae
impressed, not distinctly punctulate. Sec-
ondary sexual characters as for genus, in-
cluding (^ middle tibiae with 1 deep
excision on inner edge just before apex; $
with 2 or 3 seta-bearing punctures before
apex each side last ventral segment (punc-
tures unsymmetric in the single specimen);
9 unkno\\n. Measurements: length c. 7.7;
width c. 3.9 mm.
Type. Holotype S (Bishop Mus.) from
Bubia, Markham Vy., N-E. N. G., 50 m,
Sept. 19, 1955 (Gressitt); the type is
unique.
Notes. This species differs in form and
appearance from any other Lchia kno\\n
to me, but the generic characters, including
the excision of the i middle tibiae, are
clearly those of Lebia.
Lebia externa n. sp.
Description. With characters of genus;
form as in Figure 39; reddish yelkn\', ap-
pendages slightl)' paler; rather shining,
reticulate microsculpture absent or faint
on Iront and pronotal disc, distinct and
transverse on ebtra. Head 0.92 and 0.92
width prothorax; Iront wt'akK impressed at
middle and on each side antcriorlx , slighth'
The Carabid Beetles of New Guinea • Darlington
87
irregularly punctate. ProtJwrax rather small,
not hemispheric but transversely subquad-
rate with anterior angles broadly rounded;
width/length 1.51 and 1.52; base margined,
apex not margined at middle; disc rather
strongly transversely strigulose and vaguely
punctulate. Elytra almost 2x wide as pro-
thorax; width elytra/pro thorax 1.98 and — ;
rather strongly narrowed anteriorly; apices
obliquely truncate-emarginate with outer
angles obtuse but distinct and sutural angles
narrowly rounded; striae impressed, not
punctulate. Secondary sexual characters:
S front tarsi with squamae ( if present ) not
easily distinguishable (worn off?); S mid-
dle tibiae with 1 deep excision on inner
edge just before apex; c^ with 2, 9 4 setae
each side near apex last ventral segment.
Measurements: length 7.0-7.3; width c.
3.2-3.4 mm.
Types. Holotype $ (Bishop Mus.) from
Pinciiu, Huon Pen., N-E. N. G., Apr. 20,
1963 (Sedlacek); 1 $ paratype (M.C.Z.,
Type No. 31,399) from Wau, Morobe Dist.,
Mt. Missim, 880-1050 m, Feb. 8-9, 1963
(Sedlacek); 1 9 paratype, Popondetta,
Papua, 60 m, Sept. 3-4, 1963 (Sedlacek).
Notes. Except for the distinct outer-
apical elytral angles, this species resembles
large individuals of Lebia papiicUa, de-
scribed below.
Lebia karenia Bates
Bates 1892, Ann. Mns. Civ. Genoa 32, p. 426.
Andrewes 1933, Ent. Series Indian Forest Records
18, Part 5, pi. 3, fig. 9.
Description (of New Guinean individ-
uals). With characters of genus; form c.
of typical Lebia; head, prothorax, and lower
surface usually reddish testaceous (head
and prothorax sometimes infuscate), elytra
varying from dark with posthumeral lunules
and apices testaceous (as figin-ed by An-
drewes) to testaceous with broad sutural
anchor mark; appendages reddish or tes-
taceous; microreticulation light and irregu-
lar on front, isodiametric or slightly trans-
verse on pronotum, more transverse on
elytra. Head 0.83 and 0.79 width prothorax.
Prothorax not hemispheric but transverse-
subquadrate with anterior angles broadly
rounded; width/length 1.54 and 1.61; base
margined, apex with marginal line weak or
interrupted at middle. Elytra less than 2x
width prothorax, narrowed anteriorly; width
elytra/prothorax 1.83 and 1.84; apices
obliquely sinuate-truncate with outer and
sutural angles narrowly rounded; striae
deep, impunctate. Measurements: length
6.0-7.5; width 2.8-3.8 mm.
Types. From Burma, in Genoa Mus.
(not seen).
Occurrence in Neio Guinea. Probably
throughout New Guinea at low altitudes
and in the lower mountains up to 1200 m
(at Guega W. of Swart Valley); 20 speci-
mens seen, from all 3 political divisions of
the island.
Measured specimens. A i from Dobodura,
Papua, and 9 from TorriceHi Mts., N-E.
N. G.
Notes. My identification of this species
is based on comparison with Andrewes'
material at the British Museum.
Lebia cordifer n. sp.
Description. With characters of genus
(but 5th segments missing from all tarsi);
form (Fig. 40) of typical rather narrow
Lebia, piceous above with clypeus and
labrum, side margins of prothorax, narrow
reflexed margins of elytra, and large com-
mon heart-shaped area on elytra (extend-
ing from inside humeri to apical Vi at 2nd
intervals and reaching 6th intervals lat-
erally) testaceous; lower surface and ap-
pendages brownish to testaceous; shining,
reticulate microsculpture absent or faint
on front and on disc of pronotum (but
these areas sparsely punctulate), trans-
verse on elytra. Head 0.89 width prothorax;
eyes large and very prominent; front with
V-shaped impression at middle and im-
pressed each side anteriorly. Prothorax
relatively small, not hemispheric; width/
length 1.58; sides strongly rounded, then
strongly sinuate just before c. acute but
blunted posterior angles; base with broad,
88
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
strong, truncate lobe, weakly margined;
apex subtruncate, not margined at middle.
Elytra rather narrow but almost 2x width
of ( small ) prothorax, slightly narrowed
anteriorly; width elytra prothorax 1.95;
apices obliquely sinuate-truncate, with
outer angles broadly and sutural angles
narrowly rounded; striae impressed, not
distinctly punctulate. Secondary sexual char-
acters of i as described for genus; i with
2 setae each side before apex last ventral
segment; 9 unknown. Measurements: length
c. 5.7; width c. 2.7 mm.
Type. Holotype $ (Leiden Mus.) from
Bivak 39 A, Star Rge., West N. G., 1500 m,
July 12, 1959 (Neth. New Guinea Exp.);
the type is unique.
Notes. This is distinguished from other
New Guinean species in the preceding Key
to Species, but I do not know its real rela-
tionships.
Lebia papuella n. sp.
Description. Form of typical Lehia with
relatively small prothorax; entirely brownish
yellow, elytra sometimes with faint darker
cloud; shining, reticulate microsculpture c.
absent on front and on disc of pronotum,
present on elytral intervals as transverse
impressions not forming regular reticula-
tions. Head 0.92 and 0.88 width prothorax;
front scarcely impressed. Prothorax small,
transversely subcjuadrate; width length 1.54
and 1.56; basal and apical marginal lines
faint or interrupted at middle; disc with
anterior transverse impression subobsolete.
Elytra much wider than prothorax, nar-
rowed anteriorly; width elytra jMothorax
1.92 and 1.91; apices oblicpiely slightK-
sinuately truncate, with outer-apical angles
broadly and sutural angles more narrowly
rounded; striae impressed, not punctate.
Secondary sexual characters as for genus;
6 with apparently 2, $ 2 or more apical
ventral setae each side. Measurements:
length 4.1-5.5; width 2.0-2.7 mm.
Types. Holotype i (M.C.Z., Type No.
31,400) and 25 paratypes all from Dobo-
dura, Papua, Mar.-July 1944 (Darlington).
Additional material. Thirtv, from nu-
merous localities in all 3 political divisions
of New Guinea, from lowlands to 1700 m
(above Wau). Some of these specimens
are assigned to this species doubtfully.
Measured specimens. The i holotype and
1 9 paratype.
Notes. Lebia papuella seems closely re-
lated to a species from Queensland, Aus-
tralia, that I identify as picipennis Macleay,
but ])apuella has less pronotal microsculp-
ture and less sinuate elytra! apices than
picipennis. Similar (but not identical) un-
determined species occur in the Philippines.
Besides the type series, I have one ex-
ceptionally large 9 from Dobodura that
seems to be papuella. (Exceptional outsize
individuals occur in some other, American,
species of Lehia.) Its proportions and mea-
surements are: head 0.87 width prothorax;
prothoracic width/length 1.54; width ehtra
prothorax 1.94; length 6.5; width 3.3 mm.
My specimens (the types) were taken
by sweeping and beating undergrowth and
low foliage in rain forest.
Lehia barda n. sp.
Description. With characters of genus;
form of typical Lebia except prothorax
tending toward hemispheric; yellow, elytra
with ± distinct common dorsal plagia dark,
the dark area sometimes extending almost
to sides of elytra; lower surface and ap-
pendages yellow; whole upper surface rel-
atively dull, with deeply impressed iso-
diametric microsculpture becoming slightK
transverse on elytra. Head 0.81 and 0.81
width prothorax; front with 2 small im-
pressions anterior!). Prothorax: width
Icngtli 1.60 and 1.70; base margined; apex
not distinctly margined at middle. Elytra
narrowed anteriorb'; width elytra prothorax
1.80 and — ; apices weakly sinuate-truncate,
witli outer and sutural angles rounded;
striae deep, not distinctK' punctulate. Sec-
ondary sext(al characters as for genus; S
apparenth w itli 2, 9 3 setae each side last
ventral segment. Mcd.surcnwnls: length
4.4-5.8; width 2.2-2.8 mm.
The Carabid Beetles of New Guinea • Darlington
89
Types. Holotype i (M.C.Z., Type No.
31,401) and 2 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. Papua: 1,
between Laloki R. and Brown R., 25 m.
Mar. 16, 1956 (Gressitt); 1, Normanby Is.,
Wakainna, Sewa Bay, Nov. 21-30, 1956
(W. W. Brandt, Bishop Mns.). N-E. N. G.:
1, Busu R., E. of Lae, 100 m, Sept. 13, 1955
(Gressitt); 1, Wewak, 2-20 m, Oct. 11,
1957 (Gressitt). West N. G.: 1, Hollandia,
Apr. 1945 (Malkin, U.S.N.M.); 1, same
locality, 100 m, Ang. 24, 1955 (Gressitt).
Measured specimens. The S holotype and
1 9 paratype from Dobodura.
Notes. This species may be related to the
preceding ( papiiello ) but has the prothorax
more hemispheric, a more distinct elytral
cloud, and much heavier dorsal microsculp-
ture. It is somewhat similar also to Lehia
mehnota Chaudoir of Australia and Java
(but not New Guinea! ) but is much smaller,
with more hemispheric prothorax, and with
the dark dorsal elytral mark less defined.
Lebia insulorum n. sp.
Description. With characters of genus;
form (Fig. 41) of typical Lehia with rather
wide prothorax; bicolored, head and pro-
thorax red, elytra piceous; lower surface
red with sides of abdomen piceous; ap-
pendages reddish testaceous; shining, re-
ticulate microsculpture absent or faint on
front and pronotum, distinct and moderately
transverse on elytra. Head 0.80 and 0.79
width prothorax; front with trace of large
but indistinct (perhaps variable) V-shaped
impression. Prothorax transverse, not hemi-
spheric; width length 1.82 and 1.87; sides
broadly rounded, slightly sinuate before
slightly obtuse, blunted posterior angles;
base margined, apex weakly or not mar-
gined at middle. Ehjtra slightly narrowed
anteriorly; width elytra/ prothorax 1.72 and
1.70; apices obliquely weakly sinuate-
truncate with outer angles broadly and
sutural angles narrowly rounded; striae
deep, not distinctly punctulate. Second-
ary .sexual characters of S as described for
genus; i with 2 setae before apex each
side last ventral segment; 9 unknown.
Measurements: length c. 7.5; width c. 3.4
mm.
Types. Holotype S (Bishop Mus.) from
Nonnanby Is., Wakaiuna, Sewa Bay,
Papua, Jan. 1-8, 1957 (Gressitt); and 1 i
paratype (C.S.I.R.O., Ganberra, Australia)
from Rossel Is., SE. Papua, Oct. 1963 (W.
W. Brandt).
Notes. Although this distinct species is
placed in relation to others in the preceding
Key to Species, I do not know its real rela-
tionships.
Genus LACHNODERMA Macleay
Macleay 1873, Trans. Ent. Soc. New South Wales
2, p. 321.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1347 ( see for additional references and list
of species ) .
Jedlicka 1963, Ent. Abhandlungen 28, p. 302.
Diagnosis. Form (Fig. 42) diagnostic;
and see preceding Key to Genera of Lebiini
of New Guinea.
Description. None required here.
Type species. Lachnoderma cinctum
Macleay, of Australia.
Generic di.sfribution. SE. Asia including
India and Japan, and across the islands
to the Philippines, New Guinea, and
Australia.
Notes. I do not know how the different
species of this genus are related to each
other, and I do not know their habitats and
habits.
Lachnoderma foveolafum Sloane
Sloane 1915, Proc. Linnean Soc. New South Wales
40, p. 472.
Description. None required here: the
only species of the genus in New Guinea;
readily recognized by form (Fig. 42), color
(see Notes below), very coarse sculpture,
and pubescence; wings full; length (to apex
of elytra) c. 8 mm.
Type. From Gairns District, North
Queensland, Australia; in Sloane Goll.,
Canberra (seen).
90
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Occurrence in New Guineo. Only in
Papua: 1, Yule Is. (Van Emden Coll.,
British Miis.); 1, Port Moresby, Sept. 24,
1955 (Gressitt), in light trap; 1, Kiunga,
Flv R., Aug. 1-3, 1957 (W. W. Brandt,
Bishop Mus.); 2, Laloki, 1909, 1910 (F.
Muir, H.S.P.A.); 2, Dogura, Oct. 20-Nov.
19, 1955 (E. L. Cassidy, Bishop Mus.); 1,
Goilala, Loloipa, Owen Stanley Rge., Jan.
1-15, 1958 (W. W. Brandt, Bishop Mus.).
Notes. Sloane's (Australian) type had the
prothorax red and elytra wholly blue-black.
Some Papuan specimens are similar but
others have the sides of the prothorax
blackish and the suture more or less red.
The variation is apparently individual.
Genus SINURUS Chaudoir
Chaudoir 1869, Ann. Soc. Ent. Belgium 12, p. 129.
Jedlicka 1963, Ent. Abhandlungen 28, pp. 298,
368 ( with key to the 3 known species ) .
Dia^nosi.s. Form (Fig. 43) diagnostic;
and see preceding Key to Genera of Lehiini
of New Guinea.
Description. None required here, but
note labrum long, emarginate, 6-setose;
mentum with short tooth; ligula very wide
(or fused with paraglossae), 4-setose; 4th
hind-tarsal segments small, simply emar-
ginate; claws with c. 4 teeth; <:5 front tarsi
with 3 segments each with 2 slender
s(iuamae at apex; S 9 both with 1 seta
each side last ventral segment.
Generic dislrihnt ion. SE. Asia (Burma,
etc.) across the islands to the Philippines
and New (guinea.
Type species. Si]uinis ojxiciis C^haudoir
( below ) .
Notes. "Sintirns?" ohsctiriis Sloane, from
Sattelberg, N-E. N.G., is transferred to
MochtJicrus (cj. v. ).
Sinurus somewhat resembles but is ap-
parently not related to Co])to^lossus of Aus-
tralia.
Sinurus opacus Chaudoir
Chaudoir 1869, Ann. .Soc. Ent. Iklgiuni 12, p. 130.
Jedlicka 1963, Ent. Al)handlungen 28, p. 368.
Louwerens 1964, Ent. Tidskiift 85, p. 188.
Description (selected characters only).
With characters of genus; form as in Figure
43; dull black; not setulose (except elytral
margins very finely setulose) but entire
upper surface heavily, finely, c. isodiamet-
rically microreticulate. Head 0.75 and 0.72
width prothorax. Protliorax variable in
shape and proportions; width length 1.27
and 1.43; base/apex 1.21 and 1.11; sides
slightly (variably) angulate near middle.
Elytra: width elytra prothorax 1.65 and
1.59; striae entire, well impressed, with
long, impressed scutellar striae. Measure-
ments ( New Guinean specimens ) : length
c. 10-11; width c. 4.4-5.2 mm.
Type. From Borneo; in Oberthiir Coll.,
Paris Mus. (not seen).
Occurrence in Neio Guinea. Papua: 1,
Popondetta, 25 m. May 1966 (Shanahan-
Lippert, Bishop Mus.), in light trap. N-E.
N. G.: 1, lower Busu R., Huon Pen., Mar.
28, 1955 (E. O. Wilson, M.C.Z.), in low-
land rain forest. West N. G.: 1, Araucaria
Camp, 800 m. Mar. 1939 (Toxopeus); 1,
Mt. Gyifrie, sea level-1000 ft. (-c. 300 m).
Apr. 1939 (Cheesman, S. Australian Mus.
(sic)); 3, Waigeu Is.. Camp 1, Mt. Nok.
2500 ft. (r. 760 m). May 1938 (Cheesman).
Measured specimens. Two { i 9 ) from
^^'aigeu.
Notes. The known range of opacus is
from Perak (Malay Pen.) and perhaps
Burma to the Philippines and New
(iuinea. The 7 New Guinean specimens
\'ar\ in shape and proportions of prothorax.
I cannot separate them satisfaetoriK- from
1 from Perak and 4 from the Philippines
that I have for comparison.
The few .specimens of this species that
1 ha\(> collect(xl (in the Philippines) were,
1 think, among fermenting leaves on the
ground in rain lorest.
Genus STENOTELUS Bouchard
Bouchard 1903, Ann. .Soc. Ent. France 72, p. 171.
Jedlicka 1963, Ent. .Ahliandlnnuen 28, p. 371.
Diuii^no.sis. See k'igure 44, and Key to
Genera of Lehiini of \cu- Guinea.
The Carabid Beetles of New Guinea • Darlington
91
Description. None required here, but
note labrum rather narrow, subtruncate,
not or at most faintly emarginate, 6-setose;
ligula, 4th hind-tarsal segment, claws, and
secondary sexual characters c. as described
for Sinurus (above).
Ty))e species. Stenotelus opactis Bouc-
hard.
Generic distribution. Malay Pen.,
Greater Siinda Islands, and Philippines
(opaciis); Celebes (piceiis Louwerens 1952,
Treubia 21, p. 217); and now New Guinea
(new species described below).
Notes. The species of this genus live on
tree trunks in rain forest and are probably
nocturnal.
Stenotelus spinosus n. sp.
Description. With characters of genus;
form as in Figure 44; black, appendages
dark; upper surface not pubescent, but
elytral margins very finely short-setulose;
rather shining, reticulate microsculpture c.
isodiametric on front, somewhat transverse
on disc of pronotum, more transverse on
elytra; lower surface with sparse, irregular,
short pubescence. Head 0.88 and 0.88
width prothorax; front \\'eakly impressed
each side anteriorly. Prothorax cordate
with sides angulate before middle and
strongly sinuate posteriorly (but sinuation
less than in opacus); width length 1.44 and
1.51; base/apex 1.07 and 1.07; apex mar-
gined, base not distinctly so; side margins
strongly reflexed, each with a seta at angu-
lation and at (blunted) basal angle; disc
with usual middle line and transverse im-
pressions, and faintly transversely strigu-
lose. Elytra: width elytra/prothorax 1.67
and 1.72; humeri rounded but prominent;
outer-apical angles spined, sutural angles
acutely toothed; striae entire, moderately
impressed ( but scutellar striae faint ) ; 3rd
intervals each with 2 conspicuous seta-
bearing punctures on inner edge slightly
behind middle and near apex. Inner winfi.s
full. Lei!,s slender; 4th hind-tarsal segment
long, slender, scarcely emarginate; 5th seg-
ment with short, weak accessory setae;
claws 4-toothcd, the innermost tooth small.
Secondary sexual characters: S front tarsi
scarcely dilated but with 3 segments 2-
seriately squamulose (squamae often dis-
arranged); 6 with 1, 9 2 setae each side
near apex last ventral segment. Measure-
ments: length 7.4-8.5; width 3.1-3.7 mm.
Types. Holotype S (M.C.Z., Type No.
31,402) and 13 paratypes from lower Busu
R., Huon Pen., N-E. N. G., May 4, 1955
(E. O. Wilson), in lowland rain forest; and
additional paratypes as follows. Papua: 5,
Dobodura, Mar.-July 1944 (Darlington);
1, Kiunga, Fly R., Aug. 24-27, 1957 (W. W.
Brandt, Bishop Mus. ); 4, Normanby Is.,
Wakaiuna Bay, Dec. 1-10, 1956, and Jan.
1-8, 1957 (W. W. Brandt, Bishop Mus.).
West N. G.: 1, Mt. Nomo, S. of Mt.
Bougainville, 700 ft. (c. 210 m), Feb. 1936
(Cheesman); 2, Waigeu Is., Camp 1, Mt.
Nok, 2500 ft. {c. 760 m), May 1938 (Chees-
man ) .
Measured specimens. The S holotype and
1 9 paratype from Dobodura.
Notes. S. spinosus is probably related to
S. piceus Louwerens of Celebes (see under
Generic distriI)ution, above) but piceus is
described as pubescent, with outer-apical
angles of elytra only strongly toothed, while
spinosus is not pubescent and has these
angles spined, although the length of the
spines varies.
The few specimens of this species that I
collected were taken on trunks of standing
and fallen trees in rain forest, mostly under
burlap bands put out to trap nocturnal
Carabidae.
Genus MISCELUS Klug
King 1834, Jalirbuchern Insectenkunde 1, p. 82.
Sloane 1907, Deutsche Ent. Zeitschrift for 1907,
p. 473.
1923, Trans. Ent. Soc. London for 1923,
p. 250.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae
7, p. 1359 (see for synonymy and additional
references ) .
Andrewes 1935, Fauna British India, Coleop.,
Carabidae 2, p. 3.
Jeanne! 1942, Faune de France, Coleop. Ca-
92 Bulletin Museum of Coivpomtive Zoology, Vol. 137. No. 1
rabiques, Part 2, p. 1017 (footnote: included
in "Pericalidae").
Jedlicka 1963, Ent. Alihandhingen 28, p. 398.
Diagnosis. Form (Fig. 45) characteristic
(note form of eyes and genae, and of
elytral apices); 1 or 2 setae over each eye
(see Notes below); clypeus emarginate;
labrum long, strong!)' rounded at apex, 6-
setose, emarginate in some species but not
in others; mentum toothed; ligula truncate,
with usually 4 setae at apex, and additional
setae in 2 irregular rows posteriorly; para-
glossae longer than ligula, rounded, with-
out setae; mesosternum wide between coxae;
metasternum with longitudinal row of small
tubercles each side of middle; wings full;
4th hind-tarsal segments small, oval, weakly
emarginate; 5th segments with weak ac-
cessory setae; claws not toothed; 6 front
tarsi scarcely dilated but each with 3 seg-
ments 2-seriately squamulose below; c^ with
small patch of dense pubescence on lower
edge of front femur near base; i with 1, $
2 setae each side last ventral segment, the
inner setae in 9 distant from margin.
Descri])tion. None required here.
Type species. Miscehis javanus King.
Generic distribution. SE. Asia (includ-
ing (leylon and India) to the Philippines,
New (iuinea, and part of Cape York,
Australia.
Notes. The taxonomic position of this
remarkable genus is doubtful, but will not
be debated here. Sloane (1907) suggested
a separate tribe for it, but one of the char-
acters he stressed (the presence of only
1 seta over each eye) is inconstant within
the genus (see below), and Sloane later
(1923) doubted it tribal separation was
valid. Andrewes (1935) did give it tribal
rank.
The variation in number of setae over
each eye in this genus is remarkable, it
has been noticed before, but has not been
ade(}uately described. Some ot the species,
including the type of the genus (javanus
Klug), have only 1 seta oxer each eye
(Fig. 169), while'others have 2 (Fig. 16cS).
Many species ol Carabidae belonging to
genera that normally have 2 pairs of setae
over the eyes are known to have lost the
anterior pair, but the posterior setae then
usually remain in their original position,
between or slighth' behind the posterior
corners of the eyes. But in the Miseelus
with a single seta over each eye, the seta is
between the positions of the 2 original ones,
and appears to correspond to the single
seta over each eye of the tribe Harpalini.
The New Guinean Miseelus with 1 and with
2 setae over each eye are apparenth' dif-
ferent species, but they are so similar that
some authors ( not noticing the setae ) have
failed to separate them or have treated them
as "varieties." Intermediates do not usually
occur: each individual has either 2 setae
over each eye or 1 seta in intermediate
position. The only exception 1 have found
is a 9 unicolor from Geelvink Bay (Paris
Mus. ) with 1 seta each side in intermediate
position and also, but only on the left side,
an additional seta posteriorly. Most com-
mon species of Miseelus have 1 seta over
each eye, but forms with 2 occur in Ceylon
and southeastern Asia as well as in New-
Guinea. I plan to consider this case in
more detail in Part I\' of the present work,
in discussion of variation of taxonomic
characters.
The variation of the labrum, entire or
emarginate in different members of this
genus, is noteworthy too.
The species of Miseelus that I ha\(' col-
lected in New Guinea and the Philippines
were on or under the bark of [vcv trunks or
logs in rain barest.
Key to Species ok Miscelus of Nkm- Guinea
1 . Ehtral iiitorvals 3, 5, 7 carinati' at liaso;
( 2 setae ()\ cr each eye; prothora.v more
(juadrate; len.uth 14.5 mm ) (p. 93) lit(tuit\us
- Ehtral interxals not carinale at l)a.se 2
2. Two setae o\er eaeli eye; lalirnm with ape.\
emariiiiiate; onler-apieal el>tral angles more
narrowly rounded (p. 93) .sihliit^
- One seta o\er eacli eye; lahruni iidt em;u-
jiinate; outer-apical angles oi eKtri more
l)roadl\' rounded - -^
3. Not spotted (p. 93) tDtirolor
- Elytra with sul)apical sutural red spot (p.
94 ) ( jdvatui.s)
The Carabid Beetles of New Guinea • Darlington 93
Miscelus luctuosus Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 725.
Andrewes 19.35, Fauna British India, Coleop.,
Carabidae 2, p. 3, footnote.
Description. A large Miscelus with rela-
tively .square prothorax and with elytral
intervals 3, 5, and 7 carinate at base; length
14.5 mm; other distinguishing eharaeters in-
cluding number of supraocular setae and
emargination of labrum not noted by
Putzeys, but Andrewes specifies 2 setae
over each eye in this species.
Type. From Andai, Papua, New Guinea
(Beccari and D'Albertis, Genoa Mus.) (not
seen ) .
Occurrence in Netc Guinea. Apparently
known only from the type.
Notes. I have seen no Miscelus with
carinate elytral intervals from New Guinea,
although carinate forms do occur elsewhere.
I think the species is probably distinct.
It should be easily recognizable.
Miscelus sibling n. sp.
Description. With characters of genus;
form (Fig. 45) as usual; black, not spotted.
Head 0.83 and 0.78 width prothorax; 2
setae over each eye; labrum emarginate at
apex. Prothorax subcordate; width/length
1.24 and 1.22; base/apex 0.94 and 0.95;
basal transverse impression very deep.
Elytra: width elytra/prothorax 1.33 and
1.28; outer-apical angles narrowly rounded;
intervals not carinate at base. Win'^s full.
Secondary sexual characters as for genus.
Measurements: length 12.0-14.5; width c.
4.1^.5 mm.
Types. Holotype S ( Bishop Mus. ) and
4 paratypes (2 in M.C.Z., Type No. 31,403)
from Wan, Morobe Dist., N-E. N. G., 1100-
1200 m, dates in Sept., Oct., 1961, 1962
(holotype, 1100 m, Oct. 13, 1961) (Sedla-
ceks); and additional paratypes as follows.
Papua: 2 ( $ $ ), Dobodura, Mar.-July
1944 (Darlington); 3, Goilala, Loloipa,
Owen Stanlev Rge., (1 specimen 975 m),
Nov. 16-25, ' 1957 and Jan. 16-30, 1958
(W. W. Brandt, Bishop Mus.). N-E. N. G.:
2, Sattelberg, Huon Gulf, 1899 (Biro); 1,
same locality (British Mus.); 1, Karimui,
1080 m, July 14-15, 1963, (Sedlacek); 1,
Okapa, Aug. 6, 1965 ( Hornabrook ) . West
N. G.: 1, Tami, May 11, 1903 (Paris Mus.).
Measured specimens. The 6 holotype and
1 9 paratype from Dobodura.
Notes. This and the following species
(unicolor) are sympatric, occurring at sev-
eral of the same localities, and both occur
also in New Britain.
Miscelus unicolor Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 725.
?stijgicus Putzeys 1875, Ann. Mus. Civ. Genoa 7,
p. 726.
Sloane 1907, Deutsche Ent. Zeitschrift for 1907,
p. 474.
?morioformis Macleay 1876, Proc. Linnean Soc.
New South Wales 1, p. 168.
Sloane 1907, Deutsche Ent. Zeitschrift for 1907,
p. 474.
Description. None required here. This
insect, whatever its proper name (see dis-
cussion below), is the common, smaller,
unspotted Miscelus of New Guinea, with
1 seta over each eye; labrum not emar-
ginate; outer-apical angles of elytra broadly
rounded; length (in New Guinea) 9.5-13.0
mm.
Types. Of unicolor, from Java, should
be in Brussels Mus.; of stygicus, from
Andai, Papua, now in Genoa Mus.; of
morioformis, from Hall Sound, Papua, pre-
sumably in Macleay Mus., Sydney (none
seen ) .
Occurrence in Neiv Guinea. Common
and widely distributed: 69 specimens,
from numerous localities in all three polit-
ical divisions of New Guinea; most at low
altitudes, but reaching 1200 m at Wan.
Notes. The application of the name
unicolor to this species in New Guinea is
conventional. Without revising the whole
genus, which I cannot do, I cannot decide
the relationships of the New Guinean pop-
ulation to populations farther west, nor
can I decide the relationship of the un-
spotted populations to spotted javanus.
This species {"morioformis" ) is recorded
94 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
from Coen, halh\'a\- up tlie Cape York-
peninsula, Australia, by Sloane (1907).
(M/sce/us javanus Klug)
King 1834, Jahrbiichern Insectenkunde 1, p. S2, pi.
1, fig. 9.
Csiki 19.32, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1359 (see for many additional references and
conventional synonymy ) .
Description. None required here. If the
typical form of this species occurs in New
Guinea, it is the only spotted Mi.scehis
there. Length c. (S.. 5-1 1.0 mm.
Types. From Java; now should be in
Berlin U. Zool. Mus. (not seen).
Occurrence in Neic Guinea. Doubtful:
New Guinea has sometimes been included
in the range of this species, but the synon-
ymy is confused and old published records
are doubtful, and I have seen no specimens
from the island.
Notes. The supposed unspotted form of
javanus, unicolor Putzeys, which may or
may not really be conspecific, does occur
in New Ciuinea and is treated above.
Genus HOLCODERUS Chaudoir
(Jliaudoir 1869, Ann. Soc. Ent. Belgium 12, p. 153.
Csiki 1932, Coleop. Cat., Caral)i(lae, Harpalinae 7,
p. 1360 (see for additional references and list
of species).
Jcdlicka 1963, Knt. Ahliandlnngen 28, p. 396.
Dia!J,iU)sis. Form (of New Guinean
species) as in Figur(> 46 (but form diverse
in species outside New Guinea); color
metallic; pronotum with 1 or more strong
setae at or near each anterior angle; elytral
apices unarmed but very strongly sinuate-
emargiiiat(>; length c. 8-9 mm.
Description (selected additional char-
acters only). Not pubescent above (sparsely
so below). Uead: labruiu moderately long,
subtiiiiicatc or slightly cmarginate, 6-setose;
inculuiii toodicd; labium 4-set()se, para-
glossac distinct, longer than labium, with-
out setae. Prolhorax: pronotum with mid-
dle line coarse. Elytra: 3rd intervals with
3 or inoic punctures, anterior punctme on
outer and middle and posterior |)uncturcs
on inner edge of intervals. Inner ir//)g.s full.
Legs: 4th hind-tarsal segments scarcely
longer than wide, shallowly emarginate;
5th segments with accessory setae; claws
with c. 4 weak teeth grouped near middle.
Secondary .sexual characters: i front tarsi
with 3 segments 2-seriately squamulose
(apical squamules of 3rd segment over-
lapping but not attached to 4th segment);
2 setae each side last ventral segment in
both sexes.
Type species. Holcodertis praemorsiis
C>haudoir, of Ceylon.
Generic distriJ)ution. SE. Asia (includ-
ing (a'vIoii and India) and across the
islands to the Philippines, New Guinea,
and northern Australia.
Notes. This genus is relatively diverse in
the western part of the Malay Archipelago.
A single species group extends eastward to
New Guhiea and Australia ( see Notes under
following species ) .
Holcoderus elongatus (Saunders)
Saunders 1863, Trans. Ent. Soc. London (3) 1,
p. 466, pi. 18, fig. 5a-h (Catascoput;) .
Wallace 1863, in Saunders paper cited above, p.
460 ( Cata.sc(>))iis).
Csiki 1932, Coleop. C^at., Carabidae, Harpalinae 7,
p. 1360 (see for additional references).
Andrewes 1946, I'roc. R. Ent. Soc. London (B) 15,
p. 87.
Description. None required here. See
Figure 46, characters stated under genus,
the following Notes, and Andrewes' (1946)
detailed rcdescription. Length c. 8-9 mm.
Type. From Dorey, West N. (i., col-
lected by Wallace; typ(^ in berlin U. Zool.
Mus. (not seen).
Occurrence in New Guinea. ProbabK
throughout New Guinea: 24 spcximcMis,
from all 3 political divisions ol the island;
most from low altitudes ( including Dobo-
dura), but 2 from Wan, 1150. 1200 m, and
1 from W'aigeu Is., 2500 ft. ( r. 760 m).
Notes. The \ariabilit\- of this species (if
it is all one species) is remarkable. The
h)rm is relatively constant, hut toloi \aries
from wholK- blue or green or copper\ to
bicolored with blue or green elytra and
The Carabid Beetles of New Guinea • Darlington 95
bright copper or violet prothorax. The
punctation of the pronotal disc varies: the
disc is always closely punctate in part, but
a variable area centered near or behind
the middle is usually less punctate. And
the lateral prothoracic setae vary in number
and position : at least 1 strong seta is always
present (unless broken off) at each pos-
terior angle, at the angulation of the pro-
thoracic margin near or just before the
middle on each side, and at each anterior
angle, but some individuals have additional
lateral setae of different sizes between the
anterior and median setae, and the occur-
rence of these extra setae is sometimes
strikingly unsymmetric.
This variation makes exact definition of
the species and comparison with other
species difficult. I think, however, that all
New Guinean specimens of the genus can
be referred to elongatus, that the latter is
probably confined to New Guinea and adja-
cent small islands, and that closely related
forms occur both in the western Malay
Archipelago (e.g., gracilis Oberthi.ir) and
in tropical northeastern Australia ( coerulei-
pennis Sloane).
I do not know the habits of Holcoderus
but I suspect that elongatus may inhabit
tree tops. Tliis would account for my
failure to find the species' natural habitat.
My single specimen from Dobodura was
taken at light, but this seems to be ex-
ceptional. No other specimens are labeled
as from light traps, and the bright color
suggests partly diurnal habits. However,
Wallace (1863) says that elongatus flies at
dusk.
Genus MINUTHODES Andrewes
Andrewes 1941, Ann. Mag. Nat. Hist. (11) 7,
p. 317.
Phitia Chaudoir 1869, Ann. Soc. Ent. Belgium 12,
p. 155 (not Platia Hiibner 1820, et al) .
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1361.
Andrewes 1939, Ann. Mag. Nat. Hist. (11) 3, p.
137.
Diagnosis. Usually immediately recog-
nizable by form (head very wide but eyes
smaller than usual in tribe, prothorax usu-
ally c. 2x wider than long, and elytra short
and subquadrate), small size (4-6.5 mm),
and other characters given in the Key to
Genera of Lehiini.
Description. Form as indicated above
and in Figures 47-58; upper surface espe-
cially of elytra often (not always) with
short pubescence, and elytra often (not
always) with color patterns of many pale
lines or pale blotches. Head wide but with
relatively small eyes; antennae rather short;
2 setae over each eye; front slightly im-
pressed each side anteriorly; clypeus sub-
truncate, with 1 seta each side; labrum
rather long, irregularly rounded or subtrun-
cate anteriorly, 6-setose; mentum toothed;
ligula with 2 principal setae and 1 or more
much smaller setae; paraglossae attached to
ligula, longer, broadly rounded, without
setae. ProtJiorax very wide, scarcely lobed
at base, very broadly emarginate anteriorly,
wdth wide, depressed or slightly reflexed
lateral margins; each margin with a seta at
basal angle and at or before middle of
length; disc with usual middle line, weak
anterior transverse impression, deeper sub-
basal transverse impression. Elytra very
wide and short; humeri prominent but
rounded; apices obliquely sinuate-truncate;
striae entire; 3rd interval with 3 dorsal
pvmctures at least in some species, but these
punctures often difficult to distinguish
amid other punctation and pubescence.
Inner wings full. Lower suiiace not or not
extensively pubescent. Legs rather slender;
tarsi sparsely setose above; 4th hind-tarsal
segment weakly emarginate; 5th segment
with accessory setae; claws each with c.
3 short, weak (vestigial?) teeth. Secondary
sexual characters: 6 front tarsi slightly
dilated, with numerous narrow squamae
not arranged in 2 series; 2 setae each side
near apex last ventral segment in both
sexes; and see under ^L sexualis for special
secondary sexual characters of this species.
Type species. Platia lineella Chaudoir,
96
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
fixed by Andrewes 1939, p. 137. Andrewes
designated this species as the type of
Platia Chaudoir, and it is therefore also
the type of MinutJiodes, proposed as a new
name for preoccupied Platia.
Generic distrihuiion. Nine species on
New Guinea and neighboring small is-
lands, fewer on the Moluccas, Celebes,
New Britain, and northern Australia;
none known elsewhere.
Notes. This is a very distinct genus, con-
fined to a limited geographic area ( above ) .
The insects live wholly or chiefly on tree
trunks and fallen logs in rain forest. Al-
though they are winged, they do not often
fly to light, which suggests that they are
mainly diurnal.
The striking secondary sexual characters
of the 9 of se.xuali.s ( and of the related
hraclnjdera Chaudoir of the Moluccas) are
unique, so far as I know.
The Greek ending -odes does not indi-
cate gender, and Andrewes did not specify
gender when he proposed Minuthodes to
replace Platia. I therefore tentatively treat
the name as feminine, to make the gender
consistent with Platia.
Key k) Si'KciEs of Mi.wrnonKS ok New Guinea
1. Ehtra luarki'd witli iiuiiicrous loiiyitiuliiial
pak' liut's, soiiK'tiiiK s iiiucli intcrniptcd 2
- Elytra differently marked or not marked 5
2. Median-lateral protlioraeie setae before mid-
dle, c. '/•; of iirotlioraeic length from apex
(p. 96) . p(i))uaii(i
- Median-lateral setae near nn'ddle of pro-
thoraeie length 3
3. Elytra dnll; (length c. 6.5 mm) (p. 97)
rossi
- Elytra shining (under pubeseenee) 4
4. Smaller (c. 4.8 mm) (p. 97) _._- scdlucikoiuni
- Larger (c. 6.2 mm) (p. 97) .suhnitcn.s
5. Metallie bine blaek (p. 9,S ) tiwlallica
- Not metallie, black with or witlioni reddish
yellow spots 6
6. Elytra not plainly pubescent ( pubesci'iiee
actually present but very short, scarcely
\isil)le); 9 last \entral segment usually
with s(iuare <'.\cision at apex, and 9 hind
femur with flange or tooth near apex an-
teriorl> ; (shiin'ng black, unspottixl or 2-
or 4-spotted, Iml il spotted at least 1 pair
of s]iols elongate) scxualis
6a. Elytra not spotted, or each with a
single pale dash behind middle (Fig.
•55) (Papua) (p. 98) scxiudls s. s.
6b. Ehtra either each with a single based
dash, or 4-spotted with posterior spots
elongate ( Figs. .56, A ) ( central and
western New Guinea) (p. 99)
_- subsp. signata
- Elytra plainl\- pubescent; 9 not as de-
scribed 7
7. lilaek, not spotted; (Goodenough Is.) (p.
99 ) si\n])h'x
- Each eh'tron with 2 rather large red spots;
(mainland of New Guinea) 8
8. Smaller (4.0-5.3 nnn); elytral spots c.
regular in outline (Fig. 57) (p. 100) __ rcgularis
- Larger ( 5.5-5.8 mm ) ; elytral .spots irregu-
lar in outline (Fig. .58) (p. 100) __ irregularis
Minuthodes papuana (Sloane)
Sloane 1917, Proc. Linnean Soe. New South W'ales
42, p. 4,33 (Platia).
Agonochda lineella Sloane 1907, Deutsche Ent.
Zeitsehrift for 1907, p. 182 (not Platia lineella
Ghaudoir 1869).
Description. With characters of genus;
form as in Figure 48; black or brownish
black, appendages irregularly brown, elytra
with complex, \ariable pattern of pale lines
(Figs. 48, A); head and pronotum mod-
erately shining although closely punctate,
elytra roughened and duller, and upper sur-
face especialh' ehtra with short but distinct
pubescence. Head 0.78, 0.79, 0.81, and 0.80
width prothorax. Prothorax: width length
2.00, 2.04, 2.00, and 1.96; base apex 1.04.
1.06, 1.05, and 1.04; median-lateral setae c.
':i prothoracic length before apex. Eh/tra:
width elytra jirothorax 1.46, 1.47, 1.45, and
1.44; outer-apical angles moderateh and
sutinal angles more narrowh' rounded. Sec-
ondary sexual characters as lor genus; 9 last
ventral segment and hind lemora not modi-
fied, ^leasnrenients: li'iigth 4.4-5.2; width
2.2-2.6 mm.
Ty))e. From lierbcrlshohe, "New Pom-
crania" ( = New Britain); should be in
Deutsches Ent. Institut, Berlin (not seen).
Occurrence in Neic Guinea. Common
and widely distributed al low altitudes
throughout New (Guinea, and octnrring
also on Nornianb\ , (ioodenousj;!!. ;in(l Hos-
The Carabid Beetles of New Guinea • Darlington 97
sel Is.: 142 specimens seen in all; reaches
at least 1200 m at Wau.
Measured specimens. A S 9 from Dobo-
dura and c^ $ from Normanby Is., figures
listed in this order.
Notes. This species occurs on New
Britain as well as New Guinea, and it
apparently represents a group of species
(or subspecies?) that includes JinceUa
( Chaudoir ) of the Moluccas ( I have a
series from Morotai Is.) and qucenslandica
( Sloane ) of North Queensland, Australia ( I
have specimens from near Cairns and from
the Rocky Scrub, Cape York Pen.). The
different forms of this group are distin-
guished mainly by elytral color pattern:
linceUa has a relatively simple pattern of
3 pale lines on each elytron ( Fig. 49 ) ;
queenslandica, a complex pattern of short
lines, with 1 or 2 longer lines formed by
fusion of short ones (Fig. 50); and papuana,
a c. intennediate but very variable pattern
(Figs. 48, A). Some specimens from New
Guinea have elytral markings like those of
the type (from New Britain) as described
by Sloane.
The elytral pattern of papuana may be
genetically dimorphic at some localities
(cf. the dimorphism of markings described
for se.xualis), but the variation as a whole
is so complex that I have been unable to
analyze it satisfactorily.
Minuthodes rossi n. sp.
Description. With characters of genus;
form as in Figure 51; brownish piceous,
elytra with pattern of many short narrow
longitudinal pale lines in 3 transverse series;
head and prothorax moderately shining al-
though closely punctate, elytra roughened
and duller, and upper surface especially
elytra with short pubescence. Head 0.71
width prothorax, narrower than usual in
genus. Prothorax: width/length 1.79; base/
apex 1.21; sides irregularly broadly rounded,
almost subangulate at middle, slightly
sinuate before well defined but slightly
obtuse basal angles; median-lateral setae
near middle of prothoracic length. Elytra:
width elytra/prothorax 1.37; outer-apical
angles broadly rounded, apices subangulate
c. opposite ends 2nd intervals, sutural
angles narrowly rounded. Secondary sexual
characters of i as for genus; 9 unknown.
Measurements: length 6.5; width 3.2 mm.
Type. Holotype $ (California Acad.)
from Maffin Bay, West N. G., Sept. 1944
(E. S. Ross); the type is unique.
Notes. This seems to be a distinct species
although known from a single specimen
from a well collected lowland locality.
Minuthodes sedlacekorum n. sp.
Description. With characters of genus;
form as in Figure 52; irregular reddish
piceous with complex elytral pattern of
many short longitudinal pale lines in 3
irregular transverse series, appendages ir-
regular testaceous and brown; upper sur-
face including elytra shining although
pubescent and moderately closely punctate.
Head 0.74 width prothorax. Prothorax:
width/length 1.78; base apex 1.23; sides
broadly arcuate, slightly sinuate before well
defined posterior angles; median-lateral
setae near middle of prothoracic length.
Elytra: width elytra/prothorax 1.49; outer-
apical angles broadly rounded, sutural
angles narrowly rounded; striae coarsely
but irregularly punctate, intervals more
finely punctate. Secondary sexual char-
acters of S as described for genus; $ un-
known. Measurements: length 4.6-4.8;
width 2.3 mm.
Type. Holotype c^ (Bishop Mus.) from
Wau, Morobe Dist., N-E. N. G., 1050 m,
Sept. 16, 1961 (Sedlaceks); 1 £ paratvpe
(M.C.Z., Type No. 31,588), Pindiu, Huon
Pen., N-E. N. G., 870-1300 m, Apr. 21-22,
1963 ( Straatman ) .
Notes. More material may show that
this is a (distinct) geographic representa-
tive of the preceding species, rossi.
Minuthodes subnitens n. sp.
Description. With characters of genus;
black, elytra with pattern (Fig. 53) of
many short longitudinal pale lines in 3
98
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
irregular transverse series, appendages red-
dish testaceous; rather shining although
whole upper surface rather closely punctate
and short-pubescent. Head 0.74 width pro-
thorax; as usual in genus except labrum
broadly emarginate at apex (an individual
rather than specific character?). Protliorax:
width length 1.84; base apex 1.25; base
more lobed than usual; sides broadly arcu-
ate, sinuate before c. right posterior angles,
with median-lateral setae near middle of
length. Elytra: width elytra prothorax
1.44; outer-apical angles broadly and sutural
angles more narrowly rounded; striae im-
pressed but not more coarsely punctate
than intervals. Secondary sexual characters
of 6 as for genus; 9 imknown. Measure-
ments: length 6.2; width 2.(S mm.
Type. Holotype $ (British Mus.) from
Mt.'Baduri, Japen Is., West N. G., 1000 ft.
(305 m), Aug. 1938 (Cheesman); the type
is unique.
Notes. This may (or may not) be a
(distinct) geographic representative of the
2 preceding species, ro.ssi and sedlacekorum.
Minufhodes metallica n. sp.
Description. With characters of genus;
lorm as in Figure 47; black, elytra with
strong blue-purple reflections, appendages
dark brown; shining but short-pubescent,
head and disc of pronotum sparsely punc-
tulate, elytra rather closely punctate as
well as punctulate. Head 0.79 width pro-
thorax. Frothorax: width length 1.98; base
ajiex 1.15; sides rather strongly rounded
anteriorly, nearly straight and conxerging
posteriorly until abruptly sinuate just be-
fore c. right posterior angles; median-lat-
eral setae c. V-s of prothoracic length from
apex. Elytra: width elytra prothorax 1.41;
outer-apical angles broadly and sutural
angles more narrowly rounded; striae
obsolete. Secondary sexual characters of
6 as lor genus; 9 unknown. Measure-
ments: IcngUi 5.0; width 2.6 mm.
Type. Holotype,' ( Briti.sh Mus.) from
Kokoda, Papua. 1300 ft. (c. 400 m). Sept.
1933 ( (>hecsinan j; 1 ,^ paratope ( S. Aus-
trahan Mus.), Mt. Lamington, Papua,
1300-1500 ft. (c. 400-460 m) (McNamara).
Notes. This is the only metallic Minu-
thodes know n from New Guinea.
Minufhodes sexualis n. sp.
Description. With characters of genus;
fonn as in Figure 55; black or brownish
black, appendages dark, elytra either with-
out markings or each with 1 pale dash on
5th interval behind middle; shining, pubes-
cence of most of upper surface absent or
so short as scarcely to be visible. Head
0.88 and 0.86 width prothorax; front
sparsely punctulate. Prothorax wide but
with relatively narrow base; width length
2.03 and 2.03; base apex 0.98 and 1.00;
median-lateral setae c. % of prothoracic
length from apex; disc sparsely punctulate.
Elytra: width elytra/prothorax 1.35 and
1.34; outer-apical angles broadly rounded,
sutural angles blunted or subdenticulate
(slightly variable); striae impressed and
punctate; intervals con\ex, without distinct
reticulate microsculpture, sparsely irregu-
larly punctate or punctulate. Secondary
sexual characters: S as for genus; 9 usu-
ally with last \entral segment with con-
spicuous c. square excision at apex, and
9 always with a short ridge or blunt tooth
on anterior edge hind femur near apex.
Measurements: length 4.5-5.6; width 2.0-
2.5 mm.
Types. Holotype 9 (M.C.Z., Type Xo.
31,404) and 1 9 paratype from Dobodma,
Papua, Mar.-July 1944 (Darlington); and
additional paratopes as lollows, all from
Papua: 7, Oro Ba\ near Dobodura, Dec.
1943-Jan. 1944 (Darlnigton); 2. Kokoda-
Pitoki, 450 m. Mar. 24, U)56 (C;ressitt); 1.
Mafuhi, 4000 ft. (1220 m), Dc>c. 1933
(C:heesman); 1, "Daradac iTn," 80 km N.
Port Moresby, 500 m, St«pt. 6, 1959 ( T. C.
Maa, Bishoi:) Mus.); 1, Koitakinunui, Apr.
1, 1918 (|. T. Ziminir, Chicago Mus.);
10, Mt. Lamington, 1300-1500 It. ( r. 400-
460 m) (McNamara. S. Australian Mus.).
Measured s))ecinicns. A 6 paratNj^e from
Kokoda-Pitoki and the ? hol()t\ii(\
The Carabid Beetles of New Guinea
Darlington
99
Notes. This species is evidently closely
related to M. bradujdcra Chaudoir of the
Moluccas (described from Batjan Is. and
represented by a series from Morotai Is.
in the M.C.Z.), but sexualis lacks the
metallic tone of the elytra of J)i(icliydcm,
and the ridge or tooth of the 9 femur, not
quite apical in scxiialis, is fully apical in
hrachijdera. These 2 forms, with the "sub-
species" described below, may eventually
be considered conspecific, but I prefer to
treat the New Guinean populations as a
separate species until their interrelationships
are better understood.
The material before me suggests that
sexualis may be dimorphic in two ways.
The pale dash on the elytron is either
present or absent but never partially de-
veloped in all specimens seen, and is some-
times present or absent in different in-
dividuals from single localities, for example
in those from Oro Bay. And, although most
females have a square excision on the last
ventral segment as described, 1 of 2 fe-
males from Dobodura has the last ventral
segment only acutely emarginate.
Minufhodes sexualis signata n. subsp.
Description. As typical sexualis (above)
except for markings ( Figs. 56, A ) : elytra
each with a broad posthumeral spot and
usually also a narrow stripe behind middle
(chiefly on 5th interval but bent inward
posteriorly) reddish or yellow^ (some in-
dividuals from Wau have only the post-
humeral stripe, as noted below). Head
0.89 and 0.90 width prothorax. Prothorax:
width/length 1.96 and 2.02; base/apex 1.03
and 1.02. Elytra: width elytra prothorax
1.41 and 1.42. Secondary sexual characters
as in typical sexualis. Measurements:
length 4.;3-5.8; width 2.0-2.9 mm.
Types. Holotype 9 (M.C.Z., Type No.
31,405) and 3 paratypes from Sambeang,
Mongi Watershed, Huon Pen., N-E. N. G.,
400 m, Apr. 21, 1955 (E. O. Wilson); and
additional paratypes as follows, all from
N-E. N. G.: 1, Butala, Mongi R., Huon
Pen., Apr. 22, 1955 (Wilson, M.C.Z.); 2,
lower Busu R., Huon Pen., Apr. 22 and May
12, 1955 (Wilson, M.C.Z.), in lowland rain
forest; 2, Finschhafen, Apr. 17 and May — ,
1944 (E. S. Ross, California Acad.); 1,
Wantoat, Finisterre Rge., 4000 ft. (1220
m), Sept. 9, 1957 (Munroe & Holland,
Canadian National Coll.); 1, Lae, 10 m,
July 5, 1962 (Sedlacek); 9, Wareo, Finsch-
hafen (L. Wagner, S. Australian Mus.);
16, Simbang, Huon Gulf, 1898 (Biro).
Additional material. N-E. N. G.: 11,
Wau, Morobe Dist., altitudes from 1050 to
1200 m, dates in Jan., Mar., Aug., Sept.,
Oct., 1961-1963 (Sedlaceks). West N. G.:
42, from localities scattered from Hollandia
to the Vogelkop.
Measured specimens. A c5 paratype from
Finschhafen and the 9 holotype.
Notes. Because this species varies geo-
graphically, I have restricted the type series
to specimens from a few localities in a
comparatively small area.
The elytral markings are essentially con-
stant, with only minor variation, in all
specimens except those from Wau, of which
only 4 have typical markings, while 7 have
markings reduced to a single posthumeral
dash on each elytron (Fig. 56 A). I have
seen no intermediates between these two
patterns. Inheritance of marking in this
case, as in typical sexualis, may be simply
Mendelian.
Minufhodes simplex n. sp.
Description. With characters of genus;
form as in Figure 54; black, not marked,
appendages brown; surface shining but
short-pubescent, head and prothorax punc-
tulate, elytra more closely punctate. Head
0.79 width prothorax. Prothorax: width/
length 2.0; base apex 1.15; sides irregularly
rounded anteriorly, nearly straight and con-
verging posteriorly, abruptly sinuate just
before c. right posterior angles; median-
lateral setae c. Vs of prothoracic length from
apex. Elytra: width elytra prothorax 1.45;
outer-apical angles broadly rounded, apices
bluntly subangulate opposite ends 2nd in-
tervals, sutural angles narrowly rounded;
100
BuUetin Museum of Comparative Zoology, Vol. 137, No. 1
striae impressed, not well defined, not
specially punctate. Secandanj .sexual char-
acters of 6 unknown; of 9 normal, with-
out special characters of sexualis. Measure-
ments: length 4.7; width 2.4 mm.
Type. Holotype $ (Manson Valentine
Coll.) from Goodenough Is., Papua, Oct.
14, 1943 (W. B. Jones); the type is unique.
Notes. I do not know whether this in-
sular species is represented on New Guinea
proper.
Minufhodes regularis n. sp.
Description. With characters of genus;
black or brownish black, appendages
brownish testaceous, elytra each with c.
regular posthumeral and subapical spots
reddish yellow (Fig. 57); rather shining al-
though surface pubescent and head and
pronotum irregularly punctulate or punc-
tate and elytra more closely punctate. Head
0.81 and 0.79 width prothorax. Prothorax:
width/length 1.96 and 1.96; base/apex 1.13
and 1.18; sides rounded anteriorly, c.
straight and converging posteriorly, briefly
but often abruptly sinuate before c. right
or slightly blunted posterior angles; median-
lateral setae c. Vy. of prothoracic length
from apex. Elytra: width elytra/prothorax
1.46 and 1.44; outer-apical angles broadly
and sutural angles narrowly rounded; striae
impressed but not sharj^ly limited and not
more coarsely punctate than intervals. Sec-
ondary sexual characters as for genus. Mea-
surements: length 4.0-5.3; width 2.1-2.8
mm.
Types. Holotype $ (M.C.Z., Type No.
31,406) and 3 jiaratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. Papua: 1,
Fly R. 5 miles l)elow Palmer R., May 23-
31, 1936 (Archbold Exp., A.M.N.H.). N-E.
N. {,.: 1, Saidor, Gabumi Village, Finis-
terre Rge., July 1-21, 1958 {\\ . \\ . Brandt,
Bishop Mus.);'2, Wan, Morobe Dist., 1150,
1200 in, Sept. 7, 1961, Sept. 26-27, 1964
(Sedlaceks); 1, Swart Vy., Karubaka, 1500
m, Sept. 20, 1958 (Gressitt), in light trap;
1, W'cwak, 2-20 m. Oct. 11, 1957 (Gressitt).
West N. G.: 1, vie. Hollandia, July-Sept.
1944 (Darlington); 1, same locality, 60 m,
Nov. 26, 1954 (L. D. Brongersma, Leiden
Mus.); 1, Maffin Bay, Aug. 1944 (E. S.
Ross, California Acad.); 1, Sibil, Star Rge.,
1260 m, Aug. 24, 1959 (Leiden Mus.); 1.
mountain slope above Bernhard Camp, 100
m, Apr. 1939 (Toxopeus).
Measured .specimens. The i holotype and
1 $ paratype from Dobodura.
Notes. M. rcfiularis is apparently widely
distributed in New Guinea at moderate
altitudes.
Minufhodes irregularis n. sp.
Description. With characters of genus;
black, elytra each with 2 (posthumeral
and subapical ) irregular reddish yellow
spots (Fig. 58), antennae and palpi red-
dish testaceous, legs much darker; rather
shining although surface short-pubescent,
head punctulate at middle and strigose at
sides, pronotal disc ± punctulate, elytra
more closely punctate and in part faintly
microreticulate. Head 0.75 and 0.77 width
prothorax, as described for gt^nus except
strigose at sides and with labrum distinctly
emarginate (both specimens). Prothorax:
width length 1.97 and 1.86; base/apex 1.12
and 1.11; sides broadly rounded, converging
posteriorly, briefly sinuate before c. right
posterior angles; median-lateral setae c. 'a
(or slightly more) of prothoracic length
from apex. Elytra: width elytra prothorax
1.34 and 1.42; outer-apical angles broadly
and sutural angles narrow I\ rounded; striae
impressed but not sharj-)!) limited and not
more coarseK' punctate than intervals. Sec-
ondary sexual characters oi S normal; 9
unknown. Mca.suroncnts: length 5.5-5.8;
w idth 2.8-2.9 mm.
Types. Holotype 6 (U.S.N. M.) and 1 S
ixuatype (M.C.Z., Type No. 31,407) both
from Hollandia, West N. (;.. May 1945
(B. Malkin).
Notes. This and th(> preceding .species
{reij,tdaris) are supcrlicialK sinu'lar, but
fh(> two are s\'miiatric and diHcr in sig-
The Carabid Beetles of New Guinea • Darlington 101
nificant details, and they may not be
closely related.
Genus CAJASCOPUS Kirby
Kirby 1825, Trans. Linncan Soc. London 14, p. 94.
Wallace 1863, in Saunders, Trans. Ent. Soc. Lon-
don (3) 1, pp. 460-461 (habits).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1362 (see for additional references).
Andrewes 1937, Proc. R. Ent. Soc. London ( B )
6, pp. 187 ff. (key to species of India, etc.).
Jeannel 1949, Coleop. Carabiques de la Region
Malgache, Part 2, p. 1007 (in text).
Jedlicka 1963, Ent. Abhandlungen 28, p. 397.
Diapwsis. See Key to Genera of Lehiini
of New Guinea. In practice most Catascoptis
can be recognized by their medium to large
size (in the tribe), form (with prominent
eyes, etc.), and usually metallic coloration,
without geometric elytral markings.
Deseription (characters common to New
Guinean species of the genus, with ex-
ceptions noted). Form variable (Figs. 59-
64), slender and convex to broad and
depressed; color metallic (except in bnm-
neus), usually green, sometimes partly or
wholly blue or purple; size c. 8-22 mm;
upper surface not pubescent, more or less
shining (elytra sometimes dull), with micro-
sculpture present or absent, if present c.
isodiametric on head, somewhat transverse
on pronotum and elytra. Head with promi-
nent eyes; 2 setae over each eye; front
longitudinally impressed each side; clypeus
± emarginate, 1-setose each side; labrum
long, rounded at apex, emarginate at apex,
6-setose; antennae with 4 segments glabrous
except for tactile setae and a little pubes-
cence at apex 4th segment; mentum toothed;
ligula 4-setose, paraglossae much longer,
not setose; palpi slender. Prothorax quadrate
or subcordate; base not lobed; lateral
margins variable, each with 1 seta at base
and 1 or more near or before middle; base
with entire margin (except in dobodiira),
apex at middle not margined or weakly so;
disc with impressed middle line, deep pos-
terior transverse impression, and usually
weak (but variable) anterior transverse
impression. Elytra with humeri prominent
but rounded (humeral margins slightly
thickened in laevigatiis); apices variable, as
described for separate species, often toothed
or spined; striae entire, punctation vari-
able; 7th intervals usually and 5th some-
times raised or carinate at base; .3rd inter-
vals usually .3-punctate (2-punctate in
Jatiis), with punctures often near middle of
intervals (not on edges) but position vari-
able. Inner icings full. Lower surface with
some inconspicuous, short, sparse pubes-
cence (much more pubescence along mid-
line in tcaUacei group); last ventral seg-
ment usually slightly, broadly (variably)
emarginate in both sexes. Legs slender;
4th hind-tarsal segments small, weakly
emarginate; 5th segments with accessory
setae; claws not toothed. Secondary sexual
characters: i front tarsi slightly ( scarcely )
dilated, with 3 segments 2-seriately squamu-
lose below; i with 1, $ 2 or 3 setae each
side last ventral segment ( except i as well
as 9 with 2 or 3 setae each side in strigicol-
lis).
Type species. C. hardwickei Kirby, of
India.
Generic distribution. Represented in 3
separate tropical areas: numerous in trop-
ical Asia and the Malay Archipelago ( and
a few in tropical Australia); fewer in
tropical Africa (absent in Madagascar);
and probably represented also in tropical
South and Central America (but Jeannel
doubts whether the American species
should be included in the genus).
Notes. Although Catascopus occurs also
in Africa and probably in tropical America,
its headquarters are in tropical Asia and
the Malay Archipelago. The greatest num-
bers of species are on the Malay Pen. and
the western part of the Archipelago, but
the genus is well represented east to New
Guinea, where 14 species are now known.
Of these 14 species, elegans and smaragdulus
range from the mainland of Asia across the
islands to northern Australia; facialis, from
Asia to western New Guinea but not Aus-
tralia; and laevigatus is common to the
102 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Moluccas and New Guinea. Most other
New Guinean species of the genus are
endemic, and one group of striking species
(the icaUacei group) has probably evolved
on New Guinea and is now represented
there by at least 5 species. Only 5
Catascoptis (2 of them endemic) occur
in Australia, and they seem to represent 5
separate invasions from New Guinea. So,
tlie distribution of the genus suggests
multiple dispersal eastward across the
Malay Archipelago, with considerable spe-
ciation and some secondary radiation on
New Guinea, and minor invasions of north-
ern Australia.
All the Catascopus that I know live on
tree trunks and fallen logs in rain forest.
They are all winged, and very active. Con-
cerning their habits, Wallace (1863) says,
"The species of the yeniis Catascopus are senii-
iiocturnal in their hal)its, never flying except
at ni^ht. The species taken at Dorev (viz.,
WaUacei, W. W. S.; elongatus, W. ' W. S.
[— Ilolcodcrus]; Anwnsis, W. W. S.; amncmis,
Clniud. [= clcfi.ans]) flew against me at dusk.
The greater part of the species and individuals
I have taken have, however, been captured
under the decaying bark of fallen trees.
"As soon as the bark of a tree splits and
cracks so as to separate it from the wood,
the Cata-icnpi frecjuent it, I)ut I could scarcely
ever capture them in that position, owing to their
great activity and the force required to tear
off the bark. After a tree has lain about a year
the bark Ix'coines rotten and can be easily
broken off, and tlicii, 1)\ thi' assistance of a
net, the insects which lurk beneath it can be
more easily caplincd. 'I'lu> larger species found
in Malacca, Borneo and Singapore used fre-
(juently to be seen coursing along the surface
of some immense fallen trees, from one crack
to another, their brilliant lioihes glittering
splendidly in the sunlight.
"To capture them was by no means easy, as
they would get under the trunk where it touched
the ground, if closely pursued and no friendly
crevice was at hantl. Many an hour ha\e 1
pleasantly spent in hunting them in tiie dcMise
swampy forests of Borneo, hi Malacca and
Singapore the spice of fear and danger would be
added to the interest of the .sport, owing to the
jirobable vicinity of tigers, who might at any
moment be watching us as eagerly and with as
di-atlly a puipose as we were watching the poor
Catascopi.
"However closely pursued 1 ha\e never seen
one of these insects fly in the day time, neither
do they come out at all into the light, except
to \'isit some part of the trunk they reside in,
to which the subcortical passages do not ex-
tend. . . . The species and indi^■iduals of this
genus are much more abundant in Malacca and
Borneo than in the equally luxuriant forests of
the Molucas and New Guinea."
Key to Species of Catascopus of New Guinea
1. Elytral apices without acute teeth or spines
at or near sutural angles 2
- Elytral apices acutely toothed or spined at
or near sutural angles 4
2. Outer elytral angles rounded or \ery
obtusely angulate 3
- Outer elytral angles right or (if obtuse)
very well defined, sometimes denticulate;
length c. 10.5-13.5 mm (p. 103) ._ facialis
3. Color metallic green or blue; length c. 8.5-
10 mm (p. 103) elegans
- Color brown or bronze; length c. 12-13
mm ( see also Notes under this species )
(p. 104) Inuniieus
4. Prothorax with 2 or more lateral setae
near or before middle on each side ( if
setae broken off, positions shown by punc-
tures ) ; Unm relatixely broad and de-
pressed 5
- Prothorax with only 1 median-lateral seta
each side; form variable but often more
slender and convex 7
5. Two or 3 setae near or before middle each
side; length 17.5 mm ( see also Descrip-
tion) (p. 104) latus I
More (often 6) such setae each side ,. (•>
6. Elytral striae lightb' impressed; ehtral
margins wider than usu;il near middle;
length r. 10-11 mm (p. 104) _ laeiifJiattis
- Elytral striae deepiT; ehtnil margins less
wide; length r. 12-13 mm (.see also
l)('scri))tii»i ) (p. 105) siiliis
7. Outer ehtral angles blunt or angulate
hut not si^ined; relati\ely small species,
usu;d!> untler 1 1 mm 8
- Outer elytral angles spined; larger species,
13-21 mm {icaUacci group) 10
8. Outer el\ tr;il ;uigles rounded or obtuse;
smaller, 'r. 7.5-8.0 nun (p. 105) _. .
_ stnaragduhis
- Outer elytral angles u.sually c. right or
acute, or if obtuse, size larger 9
9. I'rothoraeic margins narrow ( almost as in
dedans); basal marginal line of pronotum
c. obsolet(>; reticulate microsculpture oii-
.solcle on disc of el\tra; length r. 9-10
nnii (p. 106) . - clohoduro
- I'rothoracic margins slightly wider; b;isal
nuirginal line ol prothorax impres.sed;
The Carabid Beetles of New Guinea • Darlington 103
reticulate microsculptuie distinct on disc
of elytra; length 8.7-9.3 mm (p. 106) . hiroi
10. Prothoracic margins moderate 11
- Prothoracic margins wider (see Descrip-
tions ) 13
1 1 . Fifth elytral inter\ als not or not much
raised near base; length c. 13-15.5 mm
(p. 107) aruensis
- Fifth as well as 7th elytral inter\'als
raised near base; usually larger 12
12. Prothorax more rjuadrate with blunter
posterior angles; head and prothorax ±
green, elytra blue-purple (note head colored
as prothorax); length c. 15-18 mm (p.
108 ) strigicollis
- Prothorax more cordate, with more acute
posterior angles; prothorax green or
cupreous, head and elytra blue-purple
(note head colored as elytra); length c.
15-20 mm ( p. 108 ) wallacei
13. Prothorax narrower (width/length c. 1.40);
head less depressed posteriorly; head dark,
pronotum green cupreous, elytra blue-
purple or blue-green; length c. 17-22
mm (p. 109) taylori
- Prothorax wider (width/length c. 1.70);
head more depressed posteriorly; head as
well as prothorax green, elytra blue-green;
length c. 20 mm (p. 109) rex
Cafascopus facialis (Wiedemann)
Wiedemann 1819, Zoologisches Magazin 1, 3, p.
165 (Corahus).
Csiki 1932, Coleop. Cat., Carabidae, Haipalinae 7,
p. 1364 (see for synonymy, "varieties," and
many additional references not concerned with
New Guinea ) .
Jedlicka 1963, Ent. Abhandlungen 28, pp. 382, 395
( "fascialis" ) .
Description ( for recognition only ) . With
characters of genus; form rather compact;
green or blue and green; elytra with outer-
apical angles well defined, apices sometimes
subangulate (variable) near suture, striae
deeply impressed and strongly punctate,
and 5th and 7th intervals raised; length (in
New Guinea) c. 10.5-13.5 mm.
Tijpe(s). From "Bengalia," in Copen-
hagen Univ. Mus. (not seen).
Occurrence in New Guinea. West N. G. :
1, Maffin Bay, Aug. 1944 (Darlington); 1,
"Dorey" (Paris Mus.). Also 1 specimen
labeled only "N. guin" ( British Mus. ) .
Notes. If my identifications are correct,
this species ranges from SE. Asia to the
Philippines, Moluccas, and (western)
New Guinea but does not reach Australia.
It is variable, and its full synonymy and
subspecies (if any) remain to be worked
out. It is rare in New Guinea and may be
confined to the western part of the island
(perhaps it has recently arrived from the
west). I found it common on Morotai Is.
in the Moluccas.
Cafascopus elegans (Weber)
Weber 1801, Observations Entomologicae, p. 45
(Elaphnis).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1363 (see for additional references and ex-
tensive synonymy ) .
Andrewes 1937, Proc. Ent. Soc. London for 1937
(B) 6, p. 189.
Van Emden 1937, Stettiner Ent. Zeitschrift 98,
p. 35 (as subsp. australasiac Hope).
Jedlicka 1963, Ent. Abhandlungen 28, pp. 380,
385.
amoenus Chaudoir 1861, Berliner Ent. Zeitschrift
5, p. 120.
obliquatus Fairmaire 1881, Le Naturaliste 3, p.
381 ( new synonymy ) .
Description ( for recognition only ) . With
characters of genus; form convex; green or
partly coppery; prothoracic margins nar-
row; elytral apices unarmed; in general
without striking characters; length c. 8.5-
10 mm.
Types. Of elegans, from Sumatra (col-
lected by Doldorf ) , present location of type
unknown; of amoenus, from Dorey, West
N. G., now in Oberthiir Coll., Paris Mus.;
of obliquatus, from New Britain, presum-
ably now in Paris Mus. (none seen).
Occurrence in New Guinea. Very com-
mon (about 200 specimens) throughout
New Guinea, chiefly at low altitudes (in-
cluding Dobodura), but reaching 1700 m
near Wan.
Notes. The range of elegans, including
its supposed subspecies and varieties
(which need further study), is from SE.
Asia to Australia, east at least to the
Philippines and Solomons.
The name obliqiuitus Fairmaire has been
104 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
overlooked by most authors, and the cita-
tion in Csiki is incorrect. The description
clearly is based on a small specimen of
the present species, which is common in
New Britain.
Cafascopus brunneus n. sp.
Description. With characters of genus;
form as in Figure 59, compact and convex
(in genus); brown, subaeneous, append-
ages brown; rather shining, reticulate micro-
sculpture faint on head, light on pronotum
and elytra. Head large, 0.97 and 0.97 width
prothorax; front irregularly sculptured and
in part sparsely punctulatc. ProtJiorax
square-cordate; width/length 1.38 and 1.37;
base/apex 1.01 and 0.96; margins moderate;
disc lightly transversely strigulose and
punctulatc. Elytra: width elytra/prothorax
c. 1.53 and 1.59 (but elytra warped so mea-
surements inexact); humeri very prominent,
almost subangulate ( narrowly rounded ) an-
teriorly; apices oblique, scarcely sinuate,
with outer angles scarcely indicated (very
l)roadly rounded) and sutural angles nar-
rowly rounded and sometimes minutely
denticulate; striae well impressed, faintly
punctulatc; no intei'vals specially elevated
at liase. Measurements: length c. 12-13;
width c. 4.4-5.0 mm.
Types. Holotype $ ( Bishop Mus. ) and
1 9 paratype from Goilala, Tapini, Owen
Stanley Rge., Papua, 975 m, Nov. 16-25,
1957; and 2 additional paratypes (M.C.Z.,
Type No. 31,408) from Goilala, Loloipa,
Owen Stanley Rge., Jan. 16-30, Feb. 1-15.
1958 ( all these sp(>cimens, W. W. Brandt ) ;
1 paratype (S. Australian Mus.), W'areo,
Finsclihafcn, N-E. N. G. (L. Wagner).
Measured specimens. The 6 holotype and
1 9 paratype from Loloipa.
Notes. This species is unusual in its
rather compact form, j)]ain brown-aeneous
color, and simple elytral apices (excejit lor
minute, variable denticles near sutural
angles). So far as I know, il is not closely
related to any previously described species.
Characters distinguishing it from other
species are given in the preceding Key.
Cafascopus lafus n. sp.
Description. With characters of genus;
form as in Figure 60, very broad, depressed;
head and pronotum dark green, elytra
purple, lower surface and appendages red-
dish black; head and pronotum shining with
reticulate microsculpture absent or faint,
elytra dull and closely microreticulate.
Head 0.87 width prothorax; front flat,
broadly irregularly impressed. Prothorax
wide-subcordate; width length 1.77; base/
apex 0.97; side margins rather narrow (in
relation to width of prothorax ) , moderately
reflexed, left with 3, right with 2 formerly-
seta-bearing punctures at and before mid-
dle. Elytra: width elytra prothorax 1.36;
humeri broad but margin not thickened
and not subangulate; margins rather nar-
row; outer-apical angles prominent, slightly
acute; apices with moderate spines c. op-
posite ends sutural striae; striae moderateh'
impressed, scarcely punctulatc; intervals
not elevated at base, punctulatc especially
along middle, 3rd with onl\' 2 dorsal punc-
tures, less than ^4 from base and near or
behind middle (position unsymmetric).
Measurements: length 17.5; width 6.3 mm.
Type. Holotype 9 (British Mus.) from
W. Tami R., Pukusan-Humboldt Bav Dist.,
West N. G., June 1937 {\\ . Stiiber); the
type is unique.
Notes. This striking and thoroughh' dis-
tinct species is snfficientK' compared with
others in the Key to Species of Catascopus
of New Guinea.
Catascopus laevigatus Saunders
SauiKlns lcS63, Trans. Kiit. Sov. Loiulon (3) 1,
p. 458, pi. 18, fi,t,^ 2a-l).
Csiki 1932, Colcop. C"at., Caiabidae, Harpalinac 7,
p. 1365 (sor for additional rcfrrcnces ).
Description ( for recognition only). With
charact(M-s of genus; wide and d(^pr(\ssed;
green, shining; el\lra with sutural angles
spined, outer-apical ehtral angles c. right;
length c. 10-1 1 mm.
Ty))es. From ''Batch ian. Ternate and
Am. Wallace," type now in ObcM'thiir
C^oll. I'aiis Mus. (not seen).
The Carabid Beetles of New Guinea • Darlington
105
Occurrence in Neiv Guinea. Twenty-four,
from numerous localities in all 3 political
divisions of New Guinea; occurs at Do-
bodura and up to 1200 m at Wau.
Notes. I have seen specimens also from
the Am Is. and from Biiru, Cerain, and
Halniahera (Jilolo) in the Moluccas. The
closely related C. laticoUis Macleay of North
Queensland (Kuranda and Atherton Table-
land, and Coen-Rocky Scrub areas) repre-
sents the species in Australia.
Caiascopus sidus n. sp.
Description. With characters of genus;
form as in Figure 61; rather wide but less
depressed than laevig,atus; green, elytra
blue purple with green humeri ( at Wau ) or
c. wholly green (Star Rge.) or c. wholly
purple (Japen Is.), lower surface and ap-
pendages dark brown; shining, reticulate
microsculpture faint on front and on disc
of pronotum, distinct on elytra. Head 0.93,
0.90, 0.92 width prothorax; front irregularly
impressed at middle, sparsely minutely
(scarcely detectably) punctulate. Prothorax
transverse-cordate with wide base; width/
length 1.57, 1.61, 1.47; base/apex 1.14, 1.13,
1.13; side margins broader and more re-
flexed than in lacvi^atus, each with c. 6
strong setae (or punctures) in anterior %;
disc almost without transverse strigulation,
faintly and sparsely (hardly detectably)
punctulate. Elytra: width elytra/prothorax
c. 1.49, 1.54, 1.57; humeri prominent but
with margins rounded (not widened and
subangulate as in laevigatiis); outer-apical
angles well defined, c. right or nearly so;
apices with short spines not quite at sutural
angles; striae well impressed, scarcely punc-
tate ( more impressed but less punctate than
in laevigatas ) ; intervals scarcely elevated at
base. Secondarij sexual characters as de-
scribed for genus. Measurements: length c.
12-13; width 4.6-5.1 mm.
Types. Holotype $ (Bishop Mus.) from
Wau, Morobe Dist., N-E. N. G., 1200 m,
Sept. 15-30, 1962 (Sedlacek); 1 i paratype,
same locality, 1250 m, Sept. 16, 1962 ( Sed-
laceks); 1 paratype, Mt. Missim (near Wau),
1600 m, Mar. 17, 1966; 7 paratypes, Wau
Ck., 1200-1500 m, Sept. 16-18, 1964 (M.
Sedlacek) (some paratypes in M.C.Z., Type
No. 31,409).
Additional material. West N. G. : 1 ^ ,
Sibil, Star Rge., 1260 m. May 16, 1959
(Leiden Mus.), at light; 1 S, Mt. Baduri,
Japen Is., 1000 ft. (305 m), Aug. 1938
( Cheesman ) .
Measured specimens. The i holotype and
the c^ S from Star Rge. and Japen Is., in
this order.
Notes. Although similar to laevigatas,
sidus is more convex, with wider and more
reflexed prothoracic margins, and other dif-
ferential characters noted in the preceding
description. The single specimens from
Star Rge. and Japen Is. differ from the
types in color of elytra (see Description,
above) but I do not wish to call them
subspecies without seeing more material.
Caiascopus smaragdulus Dejean
Dejean 1825, Species General Coleop. 1, p. 331.
Csiki 1932, Coleop. Cat., Carabidae, Haq:)alinae 7,
p. 1366 (see for additional references and
synonymy, which do not concern New Guinea).
Description (for recognition only). With
characters of genus; small, rather broad;
green or partly coppery; prothorax with
margins wider than in clegans and set off
by submarginal longitudinal swellings;
elytra with outer-apical angles rounded or
bluntly obtuse, apices each with an acute
tooth or short spine; intervals not elevated
at base; length 8 mm or less.
Type(s). From Java; now in Oberthiir
Coll., Paris Mus. (not seen).
Occurrence in New Guinea. Seventeen
specimens from localities in all 3 political
divisions of New Guinea and from Nor-
manby Is.; at low altitudes only.
Notes. C. smaragdulus ranges from the
southern corner of Asia (Burma, etc.) to
New Guinea, New Britain, and the north-
eastern corner of Australia (specimens
from the Rocky Scrub, mid-peninsular
Cape York, taken by me in June 1932).
In this species the width of the prothorax
106 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
and the development of elytral spines vary
both individually and, I think, geographi-
cally, but I do not have enough material
from outside New Guinea to define satis-
factory subspecies.
Catascopus dobodura n. sp.
Description. With characters of genus;
form ( Fig. 62 ) c. average with elytra rather
wide but convex; green, elytra sometimes
greenish castaneous or purplish laterally,
lower surface almost black, appendages
dark brown; shining, reticulate microsculp-
ture absent or nearly so on front, pronotum,
and disc of elytra, indicated toward sides
and apex of eUtra. Head 1.01 and 1.01
\\'idth prothorax; front slightly depressed
anteriorly and longitudinally impressed each
side. Prothorax quadrate-subcordate; width
length 1.44 and 1.42; base/apex 1.08 and
1.04; sides broadly rounded anteriorly with
anterior angles only a little advanced,
strongly sinuate c. % of length before right
or slightly acute basal angles; side margins
very narrow, not set off by longitudinal
swc^llings, each with usual seta at basal
angle and 1 median-lateral seta just before
middle; basal transverse impression deep
(as usual) but basal marginal line obsolete
at middle; disc with faint transverse strigae
and faint sparse punctulation. Elytra :
width elytra/prothorax — and 1.59 (elytra
of S too spread to measure); lateral mar-
gins moderate; outer-apical angles c. right
or slightly obtuse but distinct, apices each
will) spine c. opposite end 2nd interval;
striae moderately impressed, faintly punc-
tulale; intervals no! elevated at base. Mea-
surements: length e. 9-10; width c. 3.4-
3.8 mm.
Ti/i)es. llolotype i (M.C.Z., Type No.
31,410) from Oro Bay, Papua, Dec. 1943-
jaii. 1944 ( Darlington ); and 1 9 paratyjK'
Iroin Dobodura (near Oro Bay), Mar.-Julv
1944 (Darlington).
Other material. One 9 , Kiunga, Fly R.,
July 2.3-25, 1957 (\V. W. Brandt. Bishop
Mus.).
Measured specimens. The types.
Notes. It is surprising to find a new,
medium-small, green Catascopus at low
altitudes in Papua, but the species seems
clearly distinct. In form ( except that it is
a little broader) and narrow prothoracic
margins it resembles ele<ians but is im-
mediately distinguished by spined elytral
apices, reduction of microreticulation of
elytra, and in other ways. In form of elytral
apices and reduction of microreticulation
it somewhat resembles laevii!,atus but is
more slender and convex, with only 1
median-lateral pronotal seta on each side.
The virtual obliteration of the middle part
of the posterior marginal line of the pro-
notum is diagnostic of this new species.
Catascopus hiroi n. sp.
Description. With characters of genus;
form as in preceding species (dobodura):
gr(.>en or blue-green, \\'ith some coppery
color at sides of elytra especially behind
humeri, lower surface and appendages
brown or brownish black; moderately
shining, front and disc of pronotinn with
reticulate microsculpture absent or vc^rx'
light, but disc of eh'tra entirely (trans-
versely) microreticulat(\ Head 0.99 and
1.00 width prothorax; front flat, slightK'
irregularly depressed, and with (usual)
longitudinal impression each side. Pro-
tliorax quadiati'-subcordatc; \\'idth IcMigth
1.41 and 1.34; base apex 1.13 and 1.08;
sides broadly arcuate anteriorlw sometimes
laintly angulate at median-lateral seta,
sinuat(> slightly less than ^i before right or
slightly acute basal angles; lateral margins
narrow but paralleled b\' slightly swollen
ridges acccMituating the marginal channels,
each with usual seta at basal angle and 1
median-lateral seta slightK' before middle;
basal marginal line entire in all specimens.
Elytra: width eKtra inothorax 1.63 and
1.58; ontei-apical angles sharplx delined,
right or slightK acute; apices each with
short spine near but not (juite at sutural
angle; striae well im]ii(\sse(l, slightK" iiune-
The Carabid Beetles of New Guinea • Darlington
107
tulate; intervals convex, not elevated at
base, punctulate. Measurements (of types):
length 8.7-9.3; width 3.5-3.7 mm.
Types. Holotype S ( Hungarian National
Mus.) and 2 paratypes from Stephansort,
Astrolabe Bay, N-E. N. G., 1897 (Biro);
and 1 paratype, Erima, Astrolabe Bay,
1897 (Biro). (Two paratvpes now in
M.C.Z., Type No. 31,411.)
Additional material. West N. G.: 1,
Waigeu Is., Camp 1, Mt. Nok, 2500 ft. (c.
760 m ), May 1938 ( Cheesman ) . This speci-
men is a (5 larger than the types (c. 11
mm ) and \\\\h disc of pronotum more
distinctly microreticulate, but it seems
clearly referable to hiroi.
Measured speeimens. The c5 holotype and
1 9 paratype from Stephansort.
Notes. This is another medium-small,
green species presumably related to the
preceding one (dobodura) but differing
in a number of details, including entire
basal marginal line of pronotum, presence
of distinct reticulate microsculpture on disc
of elytra, and position of elytral spines,
which are closer to the suture in bird than
in dobodura.
Cafascopus wallacei group
Catascopus wallacei Saunders and its im-
mediate relatives, including all the remain-
ing New Guinean species of the genus,
treated below, fonn an apparently natural
group of large, often conspicuously colored
species characterized by having both sutural
and outer-apical elytral angles acutely
toothed or spined and by having a longi-
tudinal zone of dense, conspicuous pubes-
cence along the midline of the body, from
prosternum almost to the tip of the ab-
domen in the i but mainly on the sterna
in the $ . Otherwise the species of this
group share the characters stated under the
genus, with minor exceptions.
The uallacei group of Catascopus centers
on New Guinea, where 5 species are now
known. Most of them are sympatric: 4 of
the 5 species have been found at Wau. Of
the 5 New Guinean species, aruensis and
wallacei each reach one or more small
islands to the west (Aru Is., Waigeu, Mysol),
and aruensis reaches also New Britain,
New Ireland, and Cape York in Australia.
A sixth species of the group (chaudoiri
Castelnau) is endemic in northern Aus-
tralia.
Some species of this group vary in-
dividually in form especially of the pro-
thorax, in degree of elevation of the 5th
elvtral intervals, and in some other details.
Although I can clearly recognize only the
5 species treated below, Straneo (see refer-
ences under the species) has distinguished
others, and he may be right. A thorough
study of long series will be required to
decide this, including study of genitalic
characters, which are indicated by Straneo.
I am, incidentally, very much indebted to
Prof. Straneo for loan of paratypes of his
3 species of this group.
Catascopus aruensis Saunders
Saunders 1863, Trans. Ent. Soc. London (3) 1,
p. 458, pi. 17, fig. 5a-b.
Csiki 1932, Coleop Cat., Carabidae, Harpalinae 7,
p. 1362 (see for additional references and
partial synonymy ) .
Straneo 1943, Ann. Mus. Civ. Genoa 61, p. 302.
Jedlicka 1963, Ent. Abhandlungen 28, pp. 382,
393.
cupricoUis Chaudoir 1883, in R. Oberthiir, Coleop-
terorum Novitates 1, p. 24 (not cupricoUis
W'aterhouse 1877).
brevispinosus Sloane 1910, Proc. Linnean Soc.
New South Wales 35, pp. 398, 400 (new
synonymy ) .
aeneicollis Andrewes 1919, Ann. Mag. Nat. Hist.
(9) 3, p. 481 (new name for cupricoUis
Chaudoir ) .
Andrewes 1924, Ann. Mag. Nat. Hist. (9) 14,
p. 593.
?dalbertisi Straneo 1943, Ann. Mus. Civ. Genoa
61, p. 304.
Description. With characters of genus
and of wallacei group (above); green or
blue-green (elytra rarely purplish); pro-
thorax more square (less cordate) than in
wallacei, but somewhat variable; elytra
with outer-apical angles spined, sutural
angles with shorter spines or simply acute
108 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
(variable); 5th intervals not or not much
elevated near base (slightly variable), 7th
intervals subcarinate at base; length c. 13-
15.5 mm.
Types. Of aruensis Samiders, from "Aru
[Is.]. Wallace," now in Oberthih- Coll.,
Paris Mus. Of cupricollis Chaudoir, from
Fly R., New Guinea, now also in Oberthiir
Coll., Paris Mus. Of brevispinostis Sloane,
from Coen, Cape York, Australia, now in
Sloane Coll., Canberra. Of dalhcHisi
Straneo, holotype from Hatam, Papua, in
Genoa Mus., and allotype from Andai,
Papua, in Straneo Coll. (See Notes, below.)
Occurrence in New Guinea. Widely
distributed but much less common than
wollaeei (below): 27 specimens, from all
3 political divisions of New Guinea and
from Normanby Is.; most from low alti-
tudes but reaching 1200 m at \Vau.
Notes. Outside New Guinea this species
occurs on the Aru Is. (type locality), New-
Britain, New Ireland, and Cape York,
Australia (types of hrevispinosus) . Pos-
sibly some of the outlying populations may
hv distinguishable as subspecies.
Of the types listed above, I have seen
only those of hrevispinosus (briefly in
1957, but Dr. B. P. Moore has sent me
additional notes on them) and the allotype
of (hilhertisi ( through the kindness of Prof.
Straneo ) .
Catascopus walJocei Saunders
Saunders 1863, Trans. Ent. Soc. London (3) 1, p.
462, pi. 17, fiK. 4a-l).
Csiki 1932, Coloop. Cat., Carabidae, Harpalinac 7,
p. 1367 (see for additional references).
Straneo 1943, Ann. Mus. Civ. Cenoa 61, p. 302,
fi.u;. a.
ledlicka 1963, Ent. Ahliandlunuen 28, pp. 382,
393.
?heccarn Straneo 1943, Ann. Mus. Civ. (ienoa 61,
p. 303, fig. h.
Description. With characters of gcinis
and of ualldcei group (above); usually
brightly bicolored, with head and elytra
purple and prothorax brassy or copjKMy,
but coloration sometimes duller; prothorax
a little more subcordat(> (less square) than
in aruensis, with posterior angles more
prominent and more acute; elytra with
sutural as well as outer-apical angles spined;
5th and 7th intervals elevated near base
(slightly variable, especially the 5th);
length c. 15-20 mm.
Tijpes. Of icallacei Saunders, from "Wa-
giou [Waigeu], Dorey and Mysol"; actual
(holo)t\'pe now in Oberthiir Coll., Paris
Mus. (not seen). Of beccarii Straneo, holo-
type from Hatam, Papua, in Genoa Mus.,
and allot\'pe from Andai, Papua, in Straneo
Coll. (allotype seen).
Occurrence in New Guinea. Common
probably throughout New Guinea: 176
specimens before me (including 82 from
Dobodura ) ; most from low altitudes, but
reaching 1300 m near Wau.
Notes. This beautiful carabid is appar-
ently confined to New Guinea and zoogeo-
graphically closely associated islands in-
cluding Aru Is., Waigeu, and Mysol. It
apparently does not reach the Moluccas
proper, nor New Britain, nor Australia.
Most of the indi\iduals from Dobodura
were trapped imder strips of burlap laid
across the trunks of fallen trees in rain
forest.
Catascopus strigicoHis Straneo
Straneo 1943, .\nn. Mus. Ci\ . Cenoa 61, p. 305,
fig. c.
Description. With characters of genus
and of udllacei group, except S as well
as 9 with 2 or 3 setae each side last ventral
segment; bicolored, head and prothorax
green or slighth' copper)", ebtra purple or
(especialK- basally) bluish or greenish; pro-
thorax almost square except sides sinuate
posteriorly (as usual in group); c^Ktra with
rather short spines at outer-apical angles
and still shorter (slightl\- variable) ones
at sutural angles; 5th as well as 7th interxals
raised near bas(>; length r. 1.5-18 mm.
Types. IIolot)pe (GcMioa Mus.) and
allot\iK> (Straneo Coll.) both from .\ndai,
Papua, Aug. 1872 (D'Albertis). I have
examined the allotype, loaned b\' courtesy
of Prof. Straneo.
The Carabid Beetles of New Guinea • Darlington
109
Occurrence in New Guinea. Papua: the
types. N-E. N. G.: 21, Wau, Morobc Dist.,
altitudes from 900 to 1500 m, dates in Mar.,
Apr., May, July, Aug., Sept., Oct., Dee.,
1961-1964 (Sedlaceks); 1, Sattelberg, Huon
Gulf, 1899 (Biro); 1, Wareo, Finschhafen
(L. Wagner, S. Australian Mus.).
Notes. This seems to be a distinct species,
immediately distinguished from wallacci by
head colored like pronotum ( not like
elytra), elytral spines shorter, and other
details, and from aruensis by 5th elytral
intervals raised, size usually larger, and S
with additional apical ventral setae. It
may occur mainly in mountains rather than
in lowlands, and it may be confined to part
of eastern New Guinea, but further collect-
ing is needed to confirm these possibilities.
Cafascopus taylori n. sp.
Description. With characters of genus
and of wallacei group; form (Fig. 63) of
large wallacei; head black or very dark blue,
prothorax brassy or slightly coppery, elytra
blue purple, lower surface and appendages
dark; head and pronotum rather shining
with reticulate microsculpture absent or
weak, elytra duller with close, slightly
transverse reticulate microsculpture. Head
0.77 and 0.81 width prothorax, impressed
across base; front with usual 2 longitudinal
impressions and slightly sculptured and
irregularly punctulate posteriorly. Prothorax
quadrate-subcordate; width length 1.55 and
1.51; base/apex 1.08 and 1.10; sides broadly
arcuate anteriorly with anterior angles flat-
tened and roundly produced, very broadly
sinuate posteriorly to right or acute slightly
denticulate posterior angles; side margins
rather wide (at widest point of prothorax,
width of the flattened margin is c. Vi
width from outer edge of margin to mid-
line of pronotum ) , flattened, reflexed, each
with usual seta at basal angle and 1 before
middle; disc finely transversely strigulose,
sparsely and faintly punctulate. Elytra long,
c. as in wallacei; width elytra/prothorax
1.42 and 1.34; outer-apical angles spined,
sutural angles acutely produced or spined
(individual variation); striae well im-
pressed, faintly punctulate; 5th and 7th
intervals elevated near base. Secondary
sexual characters normal for genus; c^ with
1, 2 2 setae each side last ventral segment.
Measurements: length c. 17-22; width c.
5.8-7.1 mm.
Types. Holotype S (M.C.Z., Type No.
31,412) from Aiura, N-E. N. G., 1900 m,
July 1962 (R. W. Taylor, #2147), in rain
forest; additional paratypes as follows.
N-E. N. G.: 4, Mt. Missim, Wau, Morobe
Dist., 950-1000, 1500, 1600-2000 m, Dec.
28, 1961, Aug. 10, Sept. 21-24, 1964 ( Sedla-
ceks); 1, Eliptamin Vy., 1200-1350 m, Aug.
16-30, 1959 (W. W. Brandt, Bishop Mus.);
4, Wareo, Finschhafen (L. Wagner, S.
Australian Mus.); 1, Moife, 15 km NW. of
Okapa, 2100 m, Oct. 7-14, 1959 (T. C. Maa,
Bishop Mus.); 2, Okapa, E. Highlands,
Apr. 20, 1964 ( Hornabrook ) ; 2, 13 km
SE. Okapa, 1650-1870 m, Aug. 26, 1964
(Sedlaceks); 2, Morae, Kukukuku [Rge.],
E. Highlands, 6000 ft. (c. 1850 m). Mar. 1,
1964 (Hornabrook). West N. G.: 3, Wissel
Lakes, Arabu Camp, 1800 m, Oct. 7, 1939
(H. Boschma, Leiden Mus.); 1, Wissel
Lakes, Enarotadi, 1800-1900 m, Aug. 10,
1963 (Sedlacek).
Measured specimens. The S holotype and
9 paratype from Enarotadi.
Notes. This new species seems close to
wallacei, from which it differs mainly in its
wider prothoracic margins. The difference
is striking on comparison of specimens.
Mainly because the prothoracic margins are
wider, the present new species has a rela-
tively narrower head and wider prothorax
as shown by measurements: in a measured
6 of wallacei the head is 0.90 width pro-
thorax and the prothoracic width/length is
1.40. Also, taylori averages larger than
wallacei and usually occurs at higher alti-
tudes.
Cafascopus rex n. sp.
Description. With characters of genus
and of wallacei group; form as in Figure
64; broad with very broad prothorax; green,
110 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
front and elytra bluish green, lower surface
and appendages brownish black; head
roughened and closely microreticulate pos-
teriorly, more shining but slightly strigulose
anteriorly, pronotum and elytra duller,
closely slightly transversely microreticulate.
Head large, but only 0.78 and 0.83 width
prothorax; depressed across base, with usual
longitudinal impression each side anteriorly.
Prothorax transverse-cordate, very wide but
with relativelv narrow base; width length
1.76 and 1.67;' base apex 0.90 and 0.89; side
margins rather broad especially anteriorly
and with anterior angles flattened and ad-
vanced, each with posterior and 1 median-
lateral seta, latter slightly farther forward
than usual. Elytra: width elytra/prothorax
1.23 and 1.29; outer-apical and sutural
angles both with short spines; striae well
impressed, scarcely punctulate; intervals
convex, slightly punctulate, 7th carinate at
base, others slightly humped but not cari-
nate. Secondary sexual characters of S nor-
mal for genus; 9 unknown. Measurements:
length c. 20; width 6.7-6.8 mm.
Types, llolotype i (Bishop Mus.) from
Mokai Village, Torricelli Mts., N-E. N. G.,
750 m, Dec. 8-15, 1958 (W. W. Brandt);
and 1 6 paratype (M.C.Z., Type No.
31,413) from Kiunga, Fly R., Papua, Sept.
24-25, 1957 (W. W . Brandt).
Notes. Within the uallacei group, the
large size and very broad, cordate pro-
thorax immediately distinguish this striking
species.
Genus PERICALUS Macleay
Maclcay 1825, Annulosa Javanica, p. 15.
Csiki 1932, Coleop. C^at., Caial)i(la(', I lurpalinac 7,
p. 1368 (sfc tor synonymy and adtlitional icln-
ences ) .
Jcannci 1919, Clolrop. C^arahiciucs dc la Rruion
Malj^aclic, Part 2, p. 1007 (in text).
Jedlicka 1963, Ent. Al)liandlungen 28, p. 373.
Dia<j,nosis. Similar to Catascopus (labrum
emarginate, ligula-paraglossae similar, 4(h
hind-larsal scgineuts scarcely emarginate,
claws simple, etc. ) but eyes more abruptl\-
jiroTuinetit; cl\]icus tnmcate; elytra usually
with geometric color pattern; size usual!)
smaller.
Description. None required here.
Type species. Pericalus cicindeloides
Macleay, of Ja\a, etc.
Generic distribution. Confined to, but
widely distributed and diverse in, tropical
Asia and the Malay Archipelago, reaching
New Guinea and New Britain but not
Australia.
Notes. The genus is represented in New
Guinea by only the following species
(fii^uratus), and on New Britain by a dif-
ferent, endemic species (klapperichi Jed-
Hcka 1953, Ent. Blatter 49, p. 145).
Pericalus figurafus Chaudoir
Chandoir 1861, Berliner Ent. Zeitschrift 5, p. 124.
Description. None required here; the
form (Fig. 65) and elytral markings make
this insect unmistakable, in New Guinea;
length c. 7-8 mm.
Type. Supposedly from Gelehes, col-
lected by Wallace; now in Oberthiir Coll.,
Paris Mus. (not seen).
Occurrence in New Guinea. Common
( more than 150 specimens ) probabK
throughout the island at low altitudes, and
occurring up to 1320 m nc>ar Wau.
Notes. So far as I know, this insect has
not been found in Celebes since Wallace's
time, and it has not been recorded from the
Moluccas. I think it is possible that the
type really cam(> from New Guinea and
that the species is endemic there. It li\es
on tree trunks and reccMith' fallen logs in
rain forest.
Genus COPTODERA Dejean
Dejean 1825, Species GeTieral (Coleop. 1, p. 273.
Csiki 1932, Coleop. Cat., Carahidae, llarpalinae 7.
p. 1370 (see for adilitional references, s>non-
ymy, and list of .species).
JcaiiiKd 1949, Coleop. Carabiqnes de la Retjion
\laluache. Part 3, pp. 921. 926.
jcdiida 19(i3. i:nt. \l)liand]nn,uen 28, p. 341.
Ecliiuxhild (;liand()ir 1883, Coleopterornni Xovi-
(atcs 1, p. 21 ( n(>\v s\i)on\in>' ).
?rri(h()C(>i)t(>(lcia Lonwerens 1958, Trenhia 24, p.
255.
The Carabid Beetles of New Guinea • Darlingion
111
Diag,nosis. See Keij to Genera of Lebiini
of Neic Guinea.
Description. Form (Figs. 66-70) broad,
± depressed; upper surface not pubescent
(in New Guinean species). Head: eyes
prominent; 2 setae over each eye; front im-
pressed each side anteriorly; clypeus with 1
seta each side; hibrum variable, usually
rather long, subtruncate or slightly emar-
ginate at apex, 6-setose; antennae with 3V-2
segments glabrous; mentum without tooth;
ligula 2-setose, paraglossae attached to but
much longer than ligula, broadly rounded,
without setae at apex but with small setae
at sides. Prof/jora.v broadly subcordate (ex-
cept in <!,rossa), with base sometimes lobed,
usually not; 2 setae each side, at basal angle
and before middle; disc with usual im-
pressions. Elytra broad; humeri broadly
rounded but rather prominent; apices with
outer-apical angles rounded, sutural angles
variable; striae entire, not distinctly punc-
tate; intervals convex but none specially
elevated; 3rd intervals with 2-4 seta-bearing
punctures (if 4, near base on outer edge,
c. Vi from base on outer edge, behind
middle on inner edge, and near apex usu-
ally on inner edge), but one or both inter-
mediate punctures missing in some species.
Inner icings full. Lower surface: proster-
num usually with a little sparse pubescence,
abdomen not pubescent. Legs slender; 4th
hind-tarsal segments simply emarginate; 5th
tarsal segments with accessory setae; claws
with 3 or 4 teeth. Secondary sexual char-
acters: i front tarsi slightly dilated, with
3 segments 2-seriately squamulose, and c^
middle tarsi with 2 segments squamulose
in some (not all) species; i middle tibiae
with 1 or 2 excisions on inner edge near
apex in most species (see Notes, below);
2 setae each side apex last ventral segment
in both sexes.
Type species. Of Coptodera, C. f estiva
Dejean, of Cuba; of Ectinocliila, E. tes-
selata Chaudoir [= aurata (Macleay)], of
Australia; of Trichocoptodera, T. maculata
Louwerens, of Celebes.
Generic distribution. In a broad sense,
the genus is pan-tropical. (In Jeannel's
restricted sense, Coptodera proper is con-
fined to the Americas, and related Old
World forms are divided into several
genera. ) In the Asiatic- Australian area,
species of the genus {sensu lato) are
numerous from southeastern Asia including
Japan across the whole Malay Archipelago,
and a few occur in Australia and New
Caledonia. For further details see Notes,
below.
Notes. Jeannel (1949) divides Cop^of/era
(sensu lato) and its immediate allies into
a number of small genera based primarily
on genitallic characters. It seems to me that
in this case, as in many others, Jeannel has
carried generic splitting beyond the limit
of usefulness. I have not attempted to
check the genitallic characters, which would
require dissection of many species from
many parts of the world. But I can say that,
if Jeannel's concept of genera were applied
to the New Guinean species, I would have
to divide Coptodera into about 5 genera,
2 or 3 of which would be new. The new
names would be meaningless except to
extreme specialists, and the fine splitting
would hide the broader relationships and
geographic patterns of the group. By treat-
ing the diverse New Guinean species as
members of one genus, I emphasize what
I think is a fact, that the group is a natural
one even though the species are diverse,
and that it has a pan-tropical distribution.
The most useful taxonomic treatment in
the end may be to retain Coptodera in a
broad sense but to divide it into a reason-
able number of natural subgenera. This
should, of course, be done on a worldwide
basis, not in a local faunal work.
Certain characters do vary remarkably in
this genus. The larger New Guinean species,
which are more typical of Coptodera, have
the base of the prothorax subtruncate, some-
times slightly oblique toward the sides but
not lobed. However, in 2 smaller New
Guinean species {papueUa and wau), the
112 Bulletin Museum of Comparative Zoology. Vol. 137, No. 1
base of the prothorax does have a distinct
short basal lobe. And in the Australian
"Ectinochih" tesseluta, the base of the
prothorax is more strongly lobed. All these
species have similar, diagnostic mouthparts
( mentum without tooth, and ligiila and
paraglossae as described), and the small
New Giiinean species are transitional in
other ways: they have wider prothoracic
margins and look more like Coptodem than
Ectinochila tesseJata does, and papueUa has
elytral markings like some more-typical
Coptodera, liut both papucUa and iiau ap-
proach Ectinochila in dense dorsal micro-
sculpture, and uau has Ectinocliila-Wke
elytral markings.
The dorsal elytral punctures confirm this
relationship. The number of punctures
varies in Coptodera. The full number is 4
on each 3rd interval, placed c. as noted in
the preceding Description. This is the ar-
rangement in the type species of the genus
(C. f estiva Dejean, of Cuba) and in some
of the more or less typical New Guinean
species, e.g., cyanclla and cluta. However,
^rossa and lincolata have the 3rd intervals
3-pimctate (puncture at basal % missing),
and oxijj)tera has the 3rd intei-vals only 2-
punctate (both median punctures missing,
leaving only the subbasal and subapical
ones). But tlie EctiiiocJiila-likv New Guinean
species (papuclla and wan) and also the
Australian E. tcssclata have the 3rd intervals
4-punctate as in typical Coptodera.
The excisi(ms of the i middle tibiae also
confirm the relationship of Ectinochila to
Coptodera. The 6 middle tibiae have a
single small excision (like that in lochia)
in inner edge near apex in the Cuban type
species of Coptodera {f estiva), in most New
(iuinc^an species of the genus including the
Ectinoclula-hkc ones, and in the Australian
E. tesselata. However, 2 mm-EctinocJiila-
like New Guinean species are different:
Coptodera ox\iptera has 2 small excisions
on each i middle tibia (like Aristolehia),
and C. ornati))ennis has none.
Most Coptodera ha\(> the dorsal surface
glabrous, but "Trichocoptodera^ maculata
Louwerens of Celebes has the pronotum
sparsely pilose. Coptodera ornatipennis
Louwerens of the Moluccas seems closely
related, and a paratype of it (which I owe
to the generosity of Mr. Louwerens) has a
few inconspicuous fine hairs still on the
pronotal disc. Specimens that I assign to
this species from New Guinea seem to
lack pronotal pubescence, but the hairs
may be rulibed off ( in light-trap speci-
mens ) or be adhering invisibly to the
pronotal surface (in specimens mounted
from alcohol). However, although I have
listed Trichocoptodera as a possible syn-
onym of Coptodera, it may eventually prove
worth recognition as a separate genus or
subgenus, distinguished by i middle tibiae
without excisions and perhaps by other
characters.
The (S species of Coptodera in New-
Guinea represent 7 stocks with different,
independent geographic distributions. ( 1 )
C. grossa is endemic and without close
relatives anywhere, so far as I know. (2) C.
ornatipennis occurs in the Moluccas as well
as New Guinea, with an apparent relati\ e
on Celebes. (3) C. cyanelki represents the
flexuosa group, which ranges from SE.
Asia to Australia (the Australian species
being r/?/.sYr^///.s- Chaudoir). (4) C lincolata
ranges from Celebes to New Guinea and
New Britain, and an apparently related
species {mastersi Macleay) is in Australia.
(5) C. cluta apparently occurs from SE.
Asia to New Guinea and New Britain,
and (6) C. oxyptcra, from Celebes to New
Guinea, New Britain, and New Ireland;
these species are not representc^d in Aus-
tralia. And (7) C. papucUa and nan are
endemic to New Guinea, probably related
to each other, and less closeK' related to
"Ectinocliihr aurata of Australia.
Most ol the common CU)pt()dcr(i in Xcw
Guinea inhabit tree trunks and recentK'
lallcn logs in rain lorcst. I low ever, a lew
species ol the genus elsewhc-re li\e among
dead leaxcs on the Uronnd. and this max* be
The Carabid Beetles of New Guinea
Darlington
113
the habitat of some of the less eommon
New Guinean ones.
Key to Species of Coptodera of New Guinea
1. Prothora.x not lobed at l)ase; dorsal micro-
reticulation moderate or partly absent; larger
( usually 5 mm or more, excepting small
individuals of lincolata ) ___ 2
- Prothorax lobed at base; dorsal micro-
reticulation close, heavily impressed; smaller
(3.5-4.8 mm) 7
2. Very large (8.5-9.5 mm); form as in Figure
66, with very long mandibles and transverse
prodiorax (p. 113) grossa
- Smaller; mandibles relatively shorter; pro-
thorax it subcordate 3
3. Head and disc of pronotum not microreticu-
late; each elytron with 2 irregidar, ± trans-
verse pale blotches ( Fig. 67 ) ; c^ middle
tibiae without excisions (p. 113) oniatipennis
- Head and disc of pronotum microreticulate
( lightly so in oxijptera ) ; c5 middle tibiae
with excision(s) on inner edge near apex _ . 4
4. Elytra with sutural angles blunt, narrowly
rounded; elytra usually (not always) con-
spicuously spotted or striped with pale 5
- Elytra with sutural angles acute, often dentic-
ulate; elytra unmarked or with only a few
inconspicuous minute pale flecks 6
5. Each elytron with 3 irregular pale blotches
sometimes joined to form an irregular longi-
tudinal stripe (p. 114) cyanella
- Elytra usually with numerous, more or less
separate, longitudinal pale lines (Fig. 68)
but pale pattern somewhat variable, some-
times almost obliterated (p. 114) ___. lineolata
6. Very broad; elytral striae less impressed;
color dark without pale markings; 3rd
elytral intervals with only 2 ( subbasal and
subapical) punctures (p. 115) oxijptcra
- Less Inroad; elytral striae deeply impressed;
elytra usually with minute pale flecks; 3rd
intervals 4-punctate (p. 115) cluia
7. Front of head heavily microreticulate but
not longitudinally rugulose; elytra irregularly
2-fasciate with pale (p. 115) papucUa
- Front of head in part longitudinally rugulose
as well as microreticulate; elytra with a
large, common, irregular X-shaped pale area
(Fig. 70) (p. 116) ivau
Copfodera grossa n. sp.
Description. With characters of genus;
form as in Figure 66; very large; reddish
black, appendages brown; .shining, elytra
faintly silky or subiridescent, reticulate
microsculpture absent or faint on front and
on disc of pronotum, distinct (but lightly
impressed) and transverse on elytra. Head
0.73 and 0.72 width prothorax; mandibles
exceptionally long, nearly straight; clypeus
rounded at sides, sinuately emarginate at
middle; labrum very long, narrowed an-
teriorly, obtusely emarginate; front almost
smooth posteriorly, slightly punctate an-
teriorly, as is clypeus. Vrothorax very wide,
formed as in Figure 66; width/length 1.93
and 1.82; base/apex 1.49 and 1.44; base
not lobed; side margins narrow, each with
seta almost at basal angle and less than
Vi from apex (farther forward than usual);
basal and apical marginal lines entire; disc
almost without punctation or strigae. Elytra
wide; width elytra prothorax c. 1.67 and
1.65; apices slightly obliquely sinuate, outer-
apical angles rounded, sutural angles blunt
or subdenticulate; striae well impressed,
punctulate; intervals convex, finely and
sparsely punctulate, 3rd with subbasal and
subapical seta-bearing punctures and 1 in-
termediate puncture on inner edge behind
middle. Secondary sexual characters: as of
genus except i with squamae on front tarsi
only ( not on middle tarsi ) ; i middle tibiae
with 1 excision; i copulatory organs as in
Figure 176. Measurements: length c. 8.5-
9.5; width 4.0-4.7 mm.
Types. Holotype $ (M.C.Z., Type No.
31,414) from Dobodura, Papua, Mar.-July
1944 (Darlington). Paratypes from N-E.
N. G.: is (Bishop Mus.), Wau (Mt.
Missim), Morobe Dist., 1100 m, July 22,
1961 (Sedlaceks); 2, Karimui, 1080 m, July
13, 1963 (Sedlacek); 1, Wareo, Finschhafen
(L. Wagner, S. Australian Mus.).
Measured specimens. The S paratype from
Wau and the 9 holotype.
Notes. Although this species scarcely
looks like a Coptodera, it has the essential
characters of the genus. It is distinguished
from other New Guinean species in the
preceding Key.
Coptodera ornatipennis Louwerens
Louwerens 1962, Tijdschrift voor Ent. 105, p. 146,
fig. 9.
Description. With characters of genus;
114 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
form and markings as in Figure 67; head
and pronotum reddish testaceous or reddish
piceous, elytra dark with pale marks as
shown, but marks somewhat variable;
reticulate microsculpture absent or faint
on front and on disc of pronotum, present
(but light) and transverse on elytra. Head
0.85 and 0.86 \\idth prothorax. ProtJiurax
wide-subcordate; width length 1.71 and
1.71; base apex 1.19 and 1.12; base not
lobed; disc with a little faint sparse punc-
tulation, not pubescent but margins with a
few short hairs near anterior angles (see
following Notes). Elytra: width elytra,
prothorax 1.61 and 1.62; sutural angles
blunt; 3rd intervals with subbasal and sub-
apical seta-bearing punctures but no inter-
mediate punctures. Secondary sexual char-
acters: 6 front tarsi \\'ith .3 segments with
squamae (as usual); 6 middle tarsi with
paired squamae at apex 1st segment and
on 2nd segment; 6 middle tibiae not ex-
cised. Measurements: length c. 5.0-6.5;
width 2..3-2.9 mm.
Types. From Amboina, Moluccas; in
Louwerens Coll. (1 paratype seen).
Occurrence in Neiv Guinea. Papua: 1,
Dobodura, Mar.-July 1944 (Darlington); 1,
Kokoda, 1200 ft. (366 m), Aug. 1933
(Cheesman). N-E. N. G.: 3, Finschhafen,
Huon Pen., 10 m, Apr. 9-16, 1963 (Sedla-
cek), in mercurv vapor light trap; 1, Pindiu,
Huon Pen., 890 m, Apr. 17, 1963 (Sedlacek).
in mercury vapor light trap. West N. (i.:
1, "Neth. New Guinea" [probably vie. Hol-
landia], Nov. 10, 1944 (T. Aarons, Cali-
fornia Acad. ) .
Measured sj)ecimens. Ai from Dobodura
and 9 from Finschhafen.
Notes. This species occurs in the MoIik*-
cas (the types) as well as in New (Guinea.
and it seems closely related to "Tricho-
coptodera" nuiculata Louwerens of C'elebcs.
My Moluccan paratype of ornalipennis
actually shows vestiges ol pronotal pubes-
cence. I can see no sign of it on the Ne\\
Guinean specimens, but the latter are prob-
ably all either from alcohol or Irom light
traps. New Guinean specimens do ha\c a
few inconspicuous short setae on the pro-
thoracic margins anteriorly, but such setae
are present in C. oxyptera too, and very
short ( vestigial? ) stubs of setae are visible
at 80x in some other species of Coptodera.
I suspect that this is a ground-living
rather than arboreal species.
Copfodera cyanella Bates
Bates 1869, Ent. Monthly Mag. 6, p. 74.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1370 (see for synon>niy and additional refer-
ences ) .
Lonwerens 1956, Treuliia 23, p. 225 (Moluccas).
Description. None required here; see
preceding Key: length ± 6-7 mm.
Type(s). From New (iuiiiea, collected
by Wallace (if really from New Guinea,
presumably collected at Dorey); now in
Oberthiir Coll., Paris Mus. (not seen).
Occurrence in New Guinea. Common
throughout New Guinea and on Normanby
Is.: 181 specimens (including 81 from
Dobodura and Oro Bay); most from low-
altitudes but up to 1200 m at Waw.
Notes. This species ranges west to the
Moluccas, ("elebes, and Borneo, and east
to New Britain. It is apparently related to
C. flcxuosa Schmidt-CToebel, which occurs
from SE. Asia to the Philippines, Celebes,
etc., overlapping the range of cyanella. C.
australis Chaudoir, of eastern Australia, is
apparently a distinct but related species.
Coptodera lineolafa Bates
Rates 1869, Knt. Mdiithly ^lat,^ 6, p. 74.
Csiki 19.>2, Coleop. (^at., Carabidae, Harpalinae 7,
p. 1371 (see lors\non\ni\ and additional refer-
ences ) .
Louwerens 1956, Trenhia 23, p. 225 (Moluccas).
Description. None required lure; see
preceding Key and Figure 68; length ± .5-6
iiiin.
'iypes. I'rom New (Fuinea, "collect(>d in
numbers | jiresumabK' at Dorey] b\' Mr.
\\ allace"; [)rcsumed t\'pe now in Oberthiir
Coll., Paris Mus. (not seen).
Occurroicc i)i Xcu- Guiwi'd. Common
l^robabK' throughout N<'w <^uinea and on
I^iak and Xonnanby Is.: 231 specimens
The Carabid Beetles of New Guinea • Darlington
115
(including 66 from Dobodura and Oro
Bay); most from low altitudes, but reaching
1200-1500 m at Wau and 1400 m at Ka-
rubaka. Swart Valley.
Notes. C. lineolata ranges from Celebes
to New Guinea and New Britain, and C.
mastcrsi Macleay of eastern Australia is
closely related.
New Guinean specimens vary in size
individually (not geographically) from c.
4.5 to 6.3 mm in length. The pale elytral
marks vary individually and perhaps also
geographically, although I cannot now de-
fine useful subspecies. Individuals from
Biak Is. have the markings notably reduced,
but variably so.
Copfodera eluta Andrewes
Andrewes 1923, Trans. Ent. Soc. London for 1923,
p. 30.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1370 ( see for synonymy and additional refer-
ences ) .
Lonwerens 1956, Treubia 23, p. 225 (Molnccas).
1964, Ent. Tidskrift 85, p. 181 (Borneo).
Jedlicka 1963, Ent. Aljhandlmmen 28, pp. 342, 349,
fiH. 103.
interrupta Chaiidoir 1869, Ann. Soc. Ent. Belsinni
12, p. 194 ( not interrupta Schniidt-Coebel
1846).
Description. None needed here; see Key
to Species of Coptodera of Neic Guinea,
and following Notes; length ± 6.5 mm.
Types. Both Chaudoir and Andrewes had
this insect from several different localities,
and neither designated a type. Its selection
should await careful study of specimens
from all pertinent localities, for the species
is variable, perhaps polytypic (Jedlicka,
1963), and often misidentified.
Occurrence in New Guinea. Twenty-six
specimens from numerous localities in all
3 political di\'isions of New Guinea; most
at low altitudes, but one at 1200-1300 m
at Wau.
Notes. This species apparently ranges
from SE. Asia to the Philippines, New
Guinea, and New Britain. Most New
Guinean individuals have the elytra slightly
flecked with pale, but some are almost
unmarked. These resemble C. oxyptera in
dark color and acute sutural angles but
differ strikingly in narrower form, deep
elytral striae, 3rd intervals 4-punctate, and
6 middle tibiae with only 1 subapical
excision.
Coptodera oxyptera Chaudoir
Chaudoir 1869, Ann. Soc. Ent. Belgium 12, p. 175.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1371 (see for additional references).
Louwerens 1956, Treubia 23, p. 225 (Moluccas).
Description ( for recognition only ) . Form
broad; color black, not marked; prothorax
usually with a few short fine setae on mar-
gins near apical angles; elytra with sutural
angles acute or acutely denticulate but not
spined; 3rd intervals with only subbasal and
subapical seta-bearing punctures; S front
but not middle tarsi squamulose, and S
middle tibiae each with 2 small excisions on
inner edge near apex; length c. 5.5-7.0 mm.
Type. From Celebes (Wallace), now in
Oberthiir Coll., Paris Mus. (not seen).
Occurrence in New Guinea. Common
probably throughout New Guinea: 145
specimens (including 77 from Dobodura);
most at low altitudes but reaching at least
1300 m at Wau and 1200 at Rattan Camp,
Snow Mts.
Notes. I have not seen specimens from
Celebes, but Chaudoii's description fits the
present species, specifying (partly by ref-
erence to his description of testrastigma)
pointed-denticulate but not spined elytral
apices and presence of only subbasal and
subapical punctures of the 3rd elytral inter-
vals. For comparison with eluta, see Notes
under that species, above. C. oxyptera, as
I identify it, occurs on Celebes, the Moluc-
eas, New Guinea, New Britain, and New
Ireland. It is not represented in Australia.
Coptodera papuella n. sp.
Description. With characters of genus;
form (Fig. 69) c. as in wau (below),
Coptodera -like but with prothorax lobed
or subpedunculate at base; dull green,
margins and much of base and apex of
prothorax and margins and markings of
116 Bulletin Museum of Comparative Zoology, Vol. 137, \o. 1
elytra testaceous, the elytral markings Ijeiiig surface. The basal lobe of the prothorax
2 transverse series of longitudinal lines of is intermediate between the usual (lobe-
varying length on the intervals, appendages less ) condition in Coptodcra and the very
irregularly testaceous; entire upper surface strong lobe in E. aiirata. In this and in
with close, heavily impressed, reticulate some other characters (see Notes under
microsculpture c. isodiametric on head and Coptodcra ) the present new species and the
disc of pronotum, scarcely transverse even following one ( icau ) connect Coptodcra
on elytra. Head 0.82 and 0.81 width pro- and Ectinochila and (I think) justify re-
thorax; mandibles rather short (in genus); ducing Ectinochila to synonymy,
labrum usually weakly emarginate at apex. Coptodcra papucUa is common on trunks
Prothorax wide-subcordate; width length and large branches of standing and recently
1.71 and 1.68; base/apex 1.14 and 1.19; fallen trees in rain forest.
base lobed, subpedunculate; base and apex
not margined at middle; disc with usual Copfodero wau n. sp.
impressions and also impressed each side. Description. With characters of genus
Elytra wide; width elytra prothorax 1.70 and (except as follows) of preceding species
and 1.68; apices obliquely sinuate-truncate (papucUa); form as in Figure 70; color c.
with outer and sutural angles rounded; as in papuclla except pale marks of elytra
striae impressed, not distinctly punctulate; fused to form a broad X, with anterior
intervals moderately convex, 3rd 4-punctate arms of X extending (narrowly) to humeri
as described for genus. Secondary .sexual and posterior arms more or less connected
characters: S front tarsi very narrowly across suture. Head 0.82 and 0.83 width
squamulose, middle tarsi not squamulose; £ prothorax; front longitudinally rugose an-
middle tibiae with 1 small excision or im- teriorly especially at sides, more irregularly
pression on inner edge near apex; 2 setae rugose posteriorly. Prothorax: width length
each side last ventral segment in both 1.69 and 1.70; base/'apex 1.26 and 1.19;
sexes. Measurements: length 3.5-3.9; width disc impressed each side (as in papuclla),
1.8-2.0 mm. with an area before middle relatively
Types. Holotype i (M.C.Z., Type No. shining and transverscK' microreticulate.
31,415) and 4 paratypes from Dobodura, Elytra: width cKtra prothorax 1.65 and
Papua, Mar.-July 1944 (Darlington), and 1.67; elytra slighth' humped near base, the
56 additional paratypes from Oro Bay raised area relatively shining and with
(near Dobodura), Dec. 1943-Jan. 1944 confused microsculpture, and other dark
(Darlington). areas of elytra slightK more shining than
Additional material. Forty-five, from nu- pale areas. Measurements: length 4.4—4.8;
merous localities in all 3 political divisions width 2.0-2.3 mm.
of New Guinea; most at low altitudes but Types. Holotype £ (Bishop Mus. ) and
up to 1150 m at Wau. Because the type 12 paratypes (some in M.C.Z., T> pe Xo.
series is adequate, because I expect to 31,416) from Wau, Morobe Dist., N-E.
distribute paratypes to all museums con- N. G., altitudes from 1200 to 1500 m, dates
cemed, and because some geographic vari- in jvme, Sept., Dee.. 1961-1962 (Sedlaceks),
ation (of markings) seems to occur, I have and additional paratxpes as follows. N-E.
restricted the type series to specinKMis Irom N. (i.: 1, Kaiiiantu, 1250 ni, Jan. 8, 1965
Dobodura and Oro liay. (Sedlacek); 3, ()kai)a, dattvs in Jan.. lune.
Measured specimens. The c5 holotype antl Sept., 1961, 1965 ( llornabrook ). West
1 9 paratype from Dobodura. N. G.: 6, I^uarotadi, Wissel Lakes, 1850-
Notes. This species resembles the Aus- 1900 in. liil\ 28, 1962 (Sedlacek).
tralian Ectinochila aurata (Macleay) in Measured specimens. The i holotype and
small size and dull, liea\ily microreticulate 1 ',■ i)aral\pi' bom Wau,
The Carabid Beetles of New Guinea • Darlington
117
Notes. For distinguishing characters and
place of this species among other New
Guinean Coptodcro, see Description above.
Notes under the preceding species (pop-
tiella ) , and tlie Key to Sj)ecies of Coptodcra
of New Guinea.
MINUPHLOEUS n. gen.
Diagnosis. See Key to Genera of Lehiini
of New Guinea., Figure 71; and Notes, be-
lo^^^
Descri})tion. Form broad, depressed;
some very short pubescence present above
and below, the hairs longer at sides of pro-
thorax and elytra. Head broad; eyes rather
small but prominent; 2 setae over each eye;
cKpeus subtruncate, 1-setose each side;
labrimi subparallel, rounded at sides an-
teriorly, notched at middle, 6-setose; an-
tennae short, reaching not or not much
beyond base of prothorax, pubescent from
apex 4th segment; mandibles ordinary;
mentum strongly toothed; ligula rather nar-
row, 4-setose; paraglossae attached to ligula
but longer, rounded, not setose; palpi
slender. Frothorax wide-subcordate; sides
of disc widely, irregularly depressed but
actual margins moderate or narrow, with
numerous lateral setae; basal marginal line
entire, apical marginal line weak or inter-
nipted at middle; disc with middle line
well impressed, basal transverse impression
very deep, anterior transverse impression
almost obsolete. Elytra: humeri prominent
but rounded, strongly margined; apices
slightly obliquely sinuate-truncate, with
outer angles broadly and sutural angles
narrowly rounded; striae entire; intervals
not elevated at base, 3rd with c. 4, 5th
with 1 or 2 (near base), 7th with c. 4 or
5 apparent special seta-bearing punctures
variable in position and difficult to identify
among other punctures. Inner wings full.
Legs: tarsi sparsely pilose above; 4th hind-
tarsal segment rather small, weakly emar-
ginate; 5th segment with accessory setae;
claws with c. 5 or 6 small teeth. Secondary
sexual characters: S front tarsi very little
dilated but with 4 segments squamulose,
the squamules slender, rather numerous,
not paired; c^ middle tarsi without squamae;
S middle tibiae with small excision on inner
edge near apex; 2 setae each side near apex
last ventral segment in both sexes; i copu-
latory organs as in Figure 177.
Type species. Minuphloeus mixttis, be-
low.
Generic distribution. That of the single
known species, below.
Notes. This insect differs from Minti-
thodes in form; the labrum is notched (not
notched in Minuthodes); and the lateral
pronotal setae are more numerous. It looks
a little like some Fhilophloeus (an Aus-
tralian genus unknown in New Guinea),
but the antennal pubescence is different
(antennae pubescent from middle of 3rd
segments in Fhilophloeus, from apex of 4th
in Minuphloeus); the labrum is different
(not notched in Fhilophloeus); etc. It
slightly resembles some Coptodera, but the
mentum is toothed (not in Coptodcra), and
the ligula is 4-setose (2-setose in Coptodcra).
Minuphloeus even resembles some wide,
depressed Catascopus, but the toothed
claws, excised i middle tibiae, and other
characters differentiate it from that genus.
I am therefore forced to treat the insect as
a new monotypic genus, exact relationships
undetermined, occurring ( so far as known )
only in a small area in New Guinea.
The name of the new genus is formed
by combining the first two syllables of
Minuthodcs with the last two of Fhilo-
phloeus.
Minuphloeus mixtus n. sp.
Description. With characters of genus;
form as in Figure 71; black, shining, most
of upper surface without reticulate micro-
sculpture but extensively punctate. Head
0.84 and 0.83 width prothorax; front ir-
regularly impressed, irregularly punctate,
with short longitudinal ridge each side in-
side position of anterior supraocular setae.
Frothorax: width/length 1.82 and 1.80;
118 BuUetin Museum of Comparatwc Zoology, Vol. 137, No. 1
base/apex 1.04 and 1.04; margins with
numerous strong setae irregularly spaced in
whole length; dise finely, sparsely, irregu-
larly punctate. EJijtro: width elytra/pro-
thorax 1.51 and 1.55; striae moderately im-
pressed, closely punctate; intervals sHghtly
con\ex, sparsely punctate. Measurements:
length c. 7.0-8.5; width c. 3.3-4.0 mm.
Types. Holotype S (Bishop Mus.) and
32 paratypes (some in M.C.Z., Type No.
31,417) from Wan (including Mt. Missim,
Kunai Ck., Mt. Kaindi), Morobe Dist., N-E.
N. G., 900, 1200, 1300, 1400, 1500, 1500-
1800, 1600-2000 m, dates in Jan., Feb.,
May, June, July, Aug., Sept., Nov., Dec,
1961-1964 (Sedlaceks), and additional para-
types as follows. N-E. N. G.: 1, Moife,
15 km NW. of Okapa, 2100 m Oct. 11-13,
1959 (T. C. Maa, Bishop Mus.); 8, Okapa,
dates in Apr., Aug., Oct., 1964, Mar. 1965
(Hornabrook). West N. G.: 19, Enarotadi,
Wlssel Lakes, 1800, 1800-1900, 1850-1950
m, dates from July 19 to Aug. 4, 1962
( Sedlacek ) .
Measured specimens. The i holotype and
1 9 paratype from VVau.
Notes. See Notes under genus. The
insect looks as if it lived on tree trunks or
under bark, but its actual habitat is not
recorded.
Genus AGONOCH/LA Chaudoir
Chaudoir 1848, Bull. Soc. Nat. Moscow 21, Part 1,
p. 119.
1869, Asm. Soc. Eiit. B(.'l,<riuin 12, p. 223.
Csiki 1932, Colcop. Cat., Caiabidae, Harpalinae 7,
p. 1379 (sec for additional references and list
of species ) .
Sloane 1898, Proc. I^inncan Soc. New South Wales
23, p. 494 (in key to Australian .uencra of
Lebiini ).
Diaii,nosis. See Key to Genera of Lebiini
of New Guinea.
Description: characters common lo the
New Guinean species of the genus (Aus-
tralian species are more diverse). Form as
in Figures 72-76; small, broad, ± convex;
short-pubescent above and below, and part
or all of upper surface also closely punctate
or (at least elytra) roughened. Head: eyes
prominent but not large; 2 setae over each
eye; clypeus subtruncate, 1-setose each side;
labrum broadly rounded or subtruncate, 6-
setose; mentum toothed; ligula with 2 long
and usually 2 short setae, paraglossae c.
as long as or slightly longer than and at-
tached to ligula; palpi, especially penulti-
mate segments, short. Protliorax variable
in form (see Figs, cited); base ± arcuate
at middle but not strongly lobed; lateral
margins narrow to \\'ide, each with seta at
base and at (usually) or slightly before
middle of length; base and apex with lightly
impressed marginal lines sometimes faint or
interrupted at middle; disc with moderate
middle line and transverse impressions.
Elytra: humeri moderately prominent,
rounded; apices obliquely sinuate-truncate,
with outer angles broadh' and sutural angles
narrowly roimded or blunted; striae entire
or nearly so but usually lightly impressed,
not sharply defined; 3rd intervals apparently
usually 3- or 4-punctate, but dorsal punc-
tures difficult to identify amid other punc-
tation and pubescence. Inner icings full.
Legs slender; 4th tarsal segments weakly
emarginate; 5th segments with accessory
setae; claws with c. 4 short teeth. Secondary
sexual characters: i front tarsi slightK'
dilated, soles formed of many squamae not
arranged in 2 series ( in all species of which
S 6 in satisfactory condition are axailable);
i middle tibiae with small notch or im-
pression on innc>r edge just before apex
(except in expa)isa); 2 setae each side last
ventral segment in both sexes.
Tyj)c sj)ecies. A. guttata Chaudoir, of
southern Australia (only spc^cies mentioned
by Chaudoir in 1848).
(U'ncric distribution. Many species in
Australia; 1 Australian species also (intro-
duced?) in Now Zralaiul; 7 small species
in New Guinea, chieih in lower nioimtains.
Notes. The 7 New Guinean species that
I assign to this genus difler among them-
seKcs, but they all seem to belong to one
small group ol (he genus that may be
The Carabid Beetles of New Guinea • Darlington 119
restricted to New Guinea and the adjacent
tropical part of Australia. Described Aus-
tralian species of the group probably in-
clude Ag,o nod lil a ovalis Sloane and intricata
Sloane ( both described 1923, Proc. Linnean
Soc. New South Wales 48, p. 39), and I
have specimens representing one or more
forms of this group from North Queensland,
Australia, from rain forest on and near the
Atherton Tableland. This group of small,
pubescent species, with notched 6 middle
tibiae, jnay prove to be worth generic sepa-
ration from A'j^onochila, but the Australian
Agonochila need much more study before
division of the genus is undertaken.
Most Australian A<ionochila live on tree
trunks, especially on shaggy-trunked Eii-
ccihjptus trees. The habitat of the New
Guinean ones is not recorded but is prob-
ably in rain forest.
Key to Species of Agonochila of New Guinea
1. Elytra with pattern of many pale longitudinal
dashes in 3 irregular transverse series which
cover nearly the whole elytra (Fig. 72) 2
- Elytra differently marked or not marked 3
2. Prothorax not depressed at sides (p. 119) ____
viinuthoides
- Prothorax depressed at sides (p. 119) . ,.
duplicata
3. Prothorax not subcordate; anterior prothoracic
angles broadly rounded-in 4
- Prothorax broadly subcordate 6
4. Elytra dark with single c. regular, common,
red or testaceous area behind middle (p.
120 ) gressitti
- Elytra not marked as described 5
5. Elytra with markings varying from isolated
pale flecks to irregular X-pattern ( Figs.
74, A, B) (p. 120) variabilis
- Color entirely red, without elytral markings
( p. 120 ) nifa
6. Elytra with 2 irregular transverse pale fasciae
behind middle (Fig. 75) or with markings
expanded (Fig. 75A); length 5.3-5.7 mm
(p. 121) cxpansa
- Elytra with a large common pale area (Fig.
76) or single broad post-median fascia;
length 6.0-6.7 mm (p. 121) dorsata
Agonochila minuthoides n. sp.
Description. With characters of genus;
form as in Figure 72; irregular dark reddish
brown, elytra with complex pattern of short
pale lines in 3 irregular transverse series,
appendages testaceous; most of upper sur-
face irregularly punctate or roughened but
surface of head and pronotum shining be-
tween punctures. Head 0.81 and 0.80 width
prothorax. Prothorax transverse-quadrate;
width length 1.57 and 1.53; base/apex 1.28
and 1.28; side margins very narrow, with no
flattened areas inside margins. Elytra:
width elytra/prothorax 1.64 and 1.66. Mea-
surements: length 4..3-4.8; width 2.2-2.4
mm.
Tiipes. Holotype $ (M.C.Z., Type No.
31,418) from Didiman Ck., Lae, N-E. N. G.,
Mar. 27, 1955 (E. O. Wilson), in lowland
rain forest; 1 $ paratype, Busu R., E. of
Lae, 100 m, Sept. 14, 1955 (Gressitt); and
1 9 paratype, Sattelberg, Huon Gulf, N-E.
N. G., 1899 (Biro).
Measured specimens. The $ holotype and
9 paratype.
Notes. The color pattern and very nar-
row prothoracic margins distinguish this
species. The 3 known specimens are all
from a rather small area in northern N-E.
N. G., but it would be unsafe to assume
that the species is really so localized.
The complex color pattern of this small
lebiine is so like that of some Minuthodes
and of Coptodcro lineolata as to suggest
mimetic convergence.
Agonochila duplicata n, sp.
Description. With characters of genus;
form as in Figure 73; irregular dark reddish
brown, elytra with complex pattern of short
pale lines in 3 irregular transverse series
(much as in preceding species, minuthoides);
appendages testaceous; much of upper sur-
face irregularly punctate, but surface shin-
ing between punctures. Head 0.71 width
prothorax. Prothorax wide; width/length
1.77; base/apex 1.25; margins broadly de-
pressed. Elytra: width elytra/prothorax
1.50. Measurements: length c. 4.5; width
2.2 mm.
Type. Holotype i (Hungarian National
120 BuUetin Mitscuiu of Comparative Zoology, Vol. 137, No. 1
Mus.) from Sattelberg, Huon Gulf, N-E.
N. G., 1899 (Biro); the type is unique.
Notes. Although the individual deseribed
above is colored much like the preceding
species (miniiflioidcs) and occurs within
the range of the latter, I think it is distinct.
The wider prothoracic margins are striking,
and the greater width they give the pro-
thorax is reflected in the proportions, the
head being relatively smaller, the prothorax
wider, and elytra relatively narrower in
dupJicata than in minuthoidc.s.
Agonochila gressitti n. sp.
Description. With characters of genus;
form slender; head and prothorax red or
reddish brown, elytra slightly darker ( often
nearly black) \\'ith large, common, red or
testaceous area behind middle varying in
size and shape but always with relatively
regular margin (compared with some fol-
lowing species); appendages testaceous;
most of upper surface punctate but mod-
erately shining between punctures. Head
0.65 and 0.68 width prothorax. Frothorax:
width length 1.72 and 1.76; base/apex not
calculated (anterior angles too rounded-in
for exact measurement of apex); sides
arcuate through most of length, sometimes
faintly subangulate at inedian-lateral setae;
posterior angles obtuse, slightly blunted;
margins moderate. Elytra: width elytra/
prothorax 1.53 and 1.62. Measurements:
length 4.2-5.5; width 2.1-2.7 mm.
Types, flolotype S (Bishop Mus.) and
5 paratypes (2 in M.C.Z., Type No. 31,419)
all from Swart Vy., Karubaka, N-E. N. G.,
1500-1550 m, dates in Nov. 1958 (Gressitt).
Addidoiial material. N-E. N. G.: 2,
Adalbert Mts., Wannma, 800-1000 ni, Oct.
26, 27, ]958 (Gressitt, 1 specimen bearing
his number 3222); 1, W'um, l^pper jiniuii
Vy., 840 m, July 16, 1955 (Gressitt)."
Measured specimens. The A holotype and
1 9 paratype.
Notes. The lorm plus coloration of this
species are diagnostic, in New Ciuiiiea.
The specimens listed under Addilional ma-
terial are slightly smaller and less sharply
bicolored than the types but seem to be
conspecific.
Agonochila rufa n. sp.
Description. With characters of genus;
form c. as in va rial) ills (following species);
entirely rufous, not marked; most of upper
surface moderately punctate but head and
pronotum shining between punctures, elytra
duller; appendages testaceous. Head 0.71
and 0.69 width prothorax. Prothorax:
width length 1.64 and 1.65; base apex not
calculated ( anterior angles rounded-in ) ;
sides faintly angulate at median-lateral
setae, slightly sinuate before c. right
(slightly obtuse) basal angles; margins
rather narrow. Elytra: width elytra pro-
thorax 1.66 and 1.63. Measurements: length
c. 4.0-4.5; width 2.1-2.3 mm.
Types. Holotype 9 (Bishop Mus.) from
Bisianumu, E. of Port Moresby, Papua, 500
m, Sept. 22, 1955 (Gressitt); and paratypes
as follows. Papua: 2 9 9 (1 in M.C.Z.,
Type No. 31,420), Kokoda-Pitoki, 450 m.
Mar. 23, 24, 1956 (Gressitt); 1 i , Keparra-
Sangi, nr. Kokoda, 500 m, ^hlr. 26, 1956
(Gressitt), "Sago palm." N-E. N. G.: 1.
Wareo, Finschhafen (L. W'agner, S. Aus-
tralian Mus.).
Measured specimens. The 9 holotype and
1 9 paratype from Kokoda-Pitoki.
Notes. The plain rufous color is diagnostic
for this species in this genus in New
Guinea.
Agonochila variabilis n. sp.
Description. With characters of genus;
form as in Figure 74; irregularh- brownish
black with \arial)le elytral markings pale
( Figs. 74, A, B ) ; appendages brownish
testaceous; most oi upper surhice punctate
but shining between punctures, elytra
slightK' l(\ss shining. Head 0.70 and 0.71
width prothorax. Prothorax: width length
1.62 and 1.67; base aj^ex not ealc-ulated
(anterior angles rounded-in ); margins mod-
erate. Elytra: width elytra prothorax 1.58
and 1.63. MeasuremcnI.s: length c. 4.0-
4.5; width 2.1-2.3 mm.
The Carabid Beetles of New Guinea • Darlington 121
Types. Holotype S ( Bishop Miis. ) and
13 paratypes (some in M.C.Z., Type No.
31,421 ) all from Wissel Lakes, West N. G.,
with following additional details: holotype
and 1 9 paratype, Uiapnra, Kamo Vy.,
1530 m, Aug. 11, 15, 1955 (Gressitt); 1
paratype, Wagete, Tigi L., 1700 m, Aug. 17,
1955 (Gressitt); 10 paratypes, Enarotadi,
altitudes from 1750 to 1900 m, dates in
Aug. 1955 (Gressitt) and July, Aug. 1962
(Sedlaeek); 1, Moanemani, Kamo V., 1500
m, Aug. 13, 1962 (Sedlaeek).
Additional material. Papua: 2, Mafulu,
4000 ft. ( c. 1230 m ) , Jan. 1934 ( Cheesman ) .
N-E. N. G.: 24, Wau, Morobe Dist., alti-
tudes from 1100 to 1450 m, dates in all
months except Apr., June, Nov., 1961-1963
(Sedlaeek); 1, Mt. Mis(s)im, Morobe Dist.,
5850 ft. ( c. 1780 m ), Apr. ( Stevens, M.C.Z.).
Measured specimens. The <5 holotype and
9 paratype from Urapura.
Notes. The specimens from Wissel Lakes
vary surprisingly in elytral pattern (Figs.
cited). Of the 2 from Mafulu, 1 has mark-
ings comparable to those of the most heavily
marked Wissel Lakes individual, and the
other is even more heavily marked. The
Wau and Mt. Mis(s)im individuals are
heavily marked ( Fig. 74B ) but somewhat
variable. The variation is obviously partly
individual, but heavy markings are ap-
parently commoner in eastern than in west-
ern New Guinea.
Agonochila expansa n. sp.
Description. With characters of genus
except as noted below; form (Fig. 75)
broad, with wide-subcordate prothorax;
black, elytra with 2 irregular, interrupted
pale fasciae behind middle, the posterior
one narrower and more interrupted, the
fasciae sometimes partly joined and ex-
tended anteriorly on each elytron (Fig. 75 A);
appendages irregularly blackish brown; en-
tire upper surface punctate but moderately
shining between punctures, and head also
obliquely-longitudinally rugulose at sides
between eyes. Head 0.76 and 0.76 width
prothorax. Prothorax: width/length 1.89
and 1.80; base/apex 1.18 and 1.18; side
margins broadly depressed, with median-
lateral setae before middle of prothoracic
length. Elytra: width elytra prothorax 1.52
and 1.59. Secondary sexual characters as
for genus except i middle tibiae not ex-
cised or impressed near apex. Measure-
ments: length 5..'3-5.7; width 2.5-2.8 mm.
Types. Holotype 9 ( Bishop Mus. ) from
Finisterre Rge., Saidor, Kiambavi Village,
N-E. N. G., Aug. 1-28, 1959 (W. W.
Brandt), and paratypes as follows. N-E.
N. G.: 1 S in poor condition (M.C.Z.,
Type No. 31,422), Wau, Morobe Dist.,
1400-1500 m, Dec. 20, 1961 (Sedlaeek); 6,
Okapa, xMar. 23, Apr. 4, 1964 (Homabrook);
1 6 (with expanded markings), 11 km S.
of Mt. Hagen (town), N-E. N. G., 2000-
2300 m. May 20, 1963 (Sedlaeek).
Measured specimens. The S paratype
from Wau and the 9 holotype.
Notes. In form and markings this species
looks more like a Coptodera than an Agono-
chila, but it has the characters of the latter
genus, as here defined. The middle tibiae
lack excisions in both 6 6 listed, but this is
probably a specific ( not generic ) character,
for the follo\\'ing species (dorsata), which
seems close in most ways to the present
one, has the notch present, but weak,
Agonochila dorsata n. sp.
Description. With characters of genus;
fonn as in Figure 76; black or irregularly
reddish black, elytra either with large testa-
ceous area as figured or the pale area re-
duced to a single transverse post-median
fascia; entire upper surface closely punc-
tate, but ± shining between punctures.
Head 0.82 and 0.82 width prothorax; front
especially at sides slightly rugulose as well
as punctate. Prothorax broadly cordate;
width length 1.84 and 1.80; base apex 1.13
and 1.15; sides broadly depressed, with
median-lateral seta slightly before middle.
Ehjtra: width elytra/ prothorax 1.58 and
1.65; sutural angles better defined than
122 Bulletin Museum of Comparatwe Zoology, Vol. 137, No. 1
usual, scarcely blunted. Secondonj sexual
characters as for genus, including 6 middle
tibiae impressed on inner edge near apex.
Measurements: length 6.0-6.7; width 2.9-
3.3 mm.
Types. Holotype S (Bishop Mus.) and
11 paratvpes (some in M.C.Z., Type No.
31,423) from Kepilam, N-E. N. G., 2420-
2540 m, June 21 and 23, 1963 (Sedlacek).
Additional material N-E. N. G.: 3, 11
km S. of Mt. Hagen (town), 2000-2300 m.
May 20, 1963 (Sedlacek); 1, Edie Ck.,
Morobe Dist., 2000-2100 m, Oct. 5-10, 1963
(Sedlaceks); 1, Kainantu, 2150 m, Jan. 8,
1965 (Sedlacek).
Measured specimens. The i holotype and
1 $ paratype.
Notes. This is the largest Agonochila in
New Guinea, and it occurs at relatively
high altitudes. The form plus markings are
diagnostic. The testaceous area of the elytra
varies geographically: it is large (c. as in
Fig. 76) in the whole type series, but
reduced to a (broad) transverse fascia
(Fig. 76A) in all specimens listed under
Additional material. However, I do not
wish to make subspecies without seeing
more material from more localities.
Genus OXYODONTUS Chaudoir
C:liaucl()ir 1869, Ann. Soc. Ent. Beljj;iuni 12, p. 239.
Diagnosis. See Keij to Genera of Lebiini
of Neic Guinea; note especially form, small
size, long acute mentum tooth, rounded-
oblique elytral apices, and plainly 3-punc-
tate 3rd elytral intervals.
Descri})tion. Form as in Figure 77; part
of surface including pronotum and sides of
elytra very inconspicuously setulose. Head:
eyes prominent, 2 setae over each eye;
labrum ± rounded, 6-setose; mentum with
long, acute tooth; ligula narrow, with 2
long and 2 shorter setae; paraglossae r.
long as ligula, attached, wide, without
setae. Vrothorax with usual 2 setae each
side. Elytra formed as figured; apices
rounded-oblique; striae entire, moderately
impressed; 3rd intervals strongly 3-punctate,
with punctures c. Vi from base on outer
edge, and near middle and apex on inner
edge. Inner icings full. Legs slender; 4th
tarsal segments small, weakly emarginate;
5th segments with accessory setae; claws
with c. 3 teeth. Secondary sexual characters:
i front tarsi slightly dilated, 3 segments
with narrow squamae not in 2 regular series;
c5 middle tibiae with minute but deep
excision on inner edge just before apex; 6
with 1, 9 2 setae each side near apex last
ventral segment.
Ty})e species. O. tripunctatus Chaudoir
( below ) .
Generic distribution. That of the single
species.
Notes. The relationships of this incon-
spicuous genus are not clear.
Oxyodontus tripunctatus Chaudoir
Chaudoir 1869, Ann. Soc. Ent. Belgium 12, p. 239.
Louwerens 1956, Treubia 23, p. 226 (Moluccas).
Description. None required here. See
under genus, of which this is the only known
species, and see Figure 77; length c. 4-4.5
mm.
Types. Two specimens from Celebes,
collected by \\'allace; type now in Ober-
thiir Coll., Paris Mus. (not seen).
Occurrence in Netc Guinea. Common
probably throughout New Guinea: 78
specimens (including 53 from Dobodura),
from localities in all 3 political dixisions of
New Guinea; chiefK' at low altitudes, but
to 1300 m at Wixu.'
Notes. This species has been prc^xiousK
recorded from Celebes and the Molueeas,
and I have a series of it (or of a closcK'
related species) also from Leytc and Lu/on
in the Philippines. It is not known in New
Britain or Australia. I think it lives in
und(>rst()ry foliage in rain forest, but my
scanl\- field notes are not ch^ar about this.
Genus MOCHTHERUS Schmidt-Goebel
Scliniitlt-(;oc])C'l 1846, Fannnia Colcop, i^irniaiiiae,
p. 76.
Csiki 1932, Coleop. Cat., Carahitlac Harpalinac 7,
The Carabid Beetles of New Guinea • Darlingion 123
p. 1382 ( see for additional references, synonymy,
and list of species).
Jedlicka 1963, Ent. Abhandlun.ucn 28, p. 352.
Dkipiosis. See Key to Genera of Lehiini
of New Guinea., and note form (Fig. 78),
unarmed elytral apices, and minutely setu-
lose pronotal disc.
Description. Form c. as in Figure 78.
Head: eyes large; 2 setae over each eye;
labiiim subtruncate, not emarginate, 6-
setose; mentum weakly, usually obtusely
(variably?) toothed; ligula and paraglossae
subequal, attached, together wide, 4-setose;
palpi slender. Prothorax cordate, with usual
2 setae each side. Elytra wide, unarmed;
apices slightly obliquely sinuate-truncate;
3rd intei'vals 2-punctate, the punctures on
inner edge behind middle and near apex.
Inner wings full. Legs slender; 4th tarsal
segments small, scarcely emarginate; 5th
segments with (weak) accessory setae;
claws each with 2 long and 1 shorter tooth.
Secondary sexual characters: $ front tarsi
scarcely dilated, 3 segments with paired
squamae; i middle tibiae not excised; i
with 1, 9 2 setae near apex each side last
ventral segment.
Type species. M. angiilatiis Schmidt-
Goebel {= tetraspilotus Macleay) of SE.
Asia, etc.
Generic distribution. SE. Asia including
Japan, and across the Malay Archipelago
to the Philippines and New Guinea (not
Australia), with one species recorded (in-
troduced?) also on Christmas Is. and
Samoa.
Notes. A single common species of the
genus occurs on New Guinea.
Mochtherus obscurus (Sloane)
Sloane 1907, Deutsche Ent. Zeitschrift for 1907,
p. 183 {?Simtms).
Andrewes 1927, Ann. Mag. Nat. Hist. (9) 19, p.
110.
immacuJatus Maindron (not Redtenbacher) 1908,
Nova Guinea 5, p. 299.
Description ( for recognition only ) . With
characters of genus; form as in Figure 78;
dull black, surface minutely short-setulose;
length c. 6-7 mm.
Type. From Sattelberg, N-E. N. G.;
should be in Deutsches Entomologisches
Institut, Berlin-Dahlem (seen by An-
drewes ) .
Occurrence in New Guinea. Common
probably throughout New Guinea ( 130
specimens, including 79 from Dobodura),
most at low altitudes, but reaching 1200
m at Wau ( only 1 specimen at this altitude )
and 1530 m on the Salawaket Rge. (2 speci-
mens). Found also on Normanby Is. (2
specimens ) and Waigeu Is. ( 1 ) .
Notes. This species occurs also on New
Britain (6 specimens including 3 from
Gazelle Pen.) and New Ireland (3).
The relationship of the New Guinean
obscurus to asemus Andrewes (recorded
from the Moluccas by Louwerens 1956,
Treubia 23, p. 226) and to other species
farther west in the Malay Archipelago re-
mains to be determined. In general the
"species" seem very closely inter-allied in
this genus, and some may prove to be
geographic subspecies.
This insect lives on and under the bark
of tree trunks and recently fallen logs in
rain forest.
(Genus MOCHTHEROIDES Andrewes)
Andrewes 1923, Trans. Ent. Soc. London for 1923,
p. 50.
Jedlicka 1963, Ent. Abhandlungen 28, p. 352.
Diagnosis. See Key to Genera of Lebiini
of New Guinea.
Description. Form c. as in Figure 79.
Head: eyes moderate; 2 setae over each
eye; mandibles moderate; labrum rounded-
truncate, not (or scarcely) emarginate, 6-
setose; mentum obtusely prominent at
middle but scarcely toothed; ligula wide,
4-setose, with paraglossae not attached (ex-
cept at base), longer and narrower than
ligula. Prothorax subcordate, with very
narrow margins, each with usual 2 setae.
Ehjtra with apices obliquely sinuate-trun-
cate, unarmed; striae entire; 3rd intervals
with 1 seta-bearing puncture on inner edge
at extreme apex but othei-wise impunctate.
124 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Legs slender; 4th tarsal segments emar-
ginate for less than half of length; 5th
segments with accessory setae; claws each
with c. 4 very small teeth. Inner icings
full. Sceonckmj sexual charoeters: 6 front
tarsi wider than in Moehfhcnts, with 3 seg-
ments 2-seriately squamulose; c5 middle
tibiae not excised; i with 1, 9 with 2
setae near apex each side last ventral seg-
ment.
Type species. Masoreiis sericans Schmidt-
Goebel, of Burma, etc.
Generic distribution. Known from
Burma, Singapore, Sumatra, Philip-
pines, and New Britain; not recorded from
New Guinea, but may occur there.
Notes. Mochthcroides superficially re-
sembles MochtJierus but the two genera are
probably not related. They differ in mouth-
parts, punctures of 3rd elytral intervals,
claw teeth, etc.
(Mochtheroides niger Jedlicka)
[t'dlic'ka 1934, Acta Soc. Ent. Prai^ue 31, p. 122.
'■ 1963, Ent. Abhandlungfii 28, p. 352.
Description ( for recognition only ) . With
characters of genus; form as in Figure 79;
black, most of surface (except prosternum)
not setulose. ProtJwrax with margins very
narrow. Elytra with 3rd intervals with only
1 (apical) seta-bearing puncture; length c.
4.5-4.8 mm.
Type. From Sibu>'an Is., Philippines;
in Andrewes Coll., British Mus. (se(Mi).
Occurrence in Netc Guinea. Not yet
found, but may occur.
Notes. Three specimens that T collected
at Cape Clouccster, New Britain (under
th(^ bark of a small dead tree) seem in-
distinguishable from the Philippine type,
with which I compared them in 194S. This
distribution suggests that the species will
be louud in New Guinea too.
Genus DOUCHOCTIS Schmidt-Goebel
Schmidt-Goebel 1846, Faunula Cok-op. Bimiainac,
p. 62.
Andrewes 1931, Zoo). Mcdedelingcn 14, pp. 62-64
(key to SuiiKitraii species).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1383 (see for additional references and list
of species ) .
Louwerens 1958, Treubia 24. pp. 258, 259 (com-
ments on some species ) .
Jedlicka 1963, Ent. Abhandlunsen 28, p. 356.
Diagnosis. Small Lebiini, rather diverse
in form; not pubescent; mentum without
tooth; ligula and paraglossae fused into a
broadly rounded whole, with usually 2
principal and several slightly smaller setae;
each 3rd elytral interval usually \N'ith 2
minute punctures behind middle, these
punctures without setae.
Description. Form variable (Figs. 80-
85); not pubescent above, with or without
reticulate microsculptme, latter ( if present)
c. isodiametric on front, transverse on
pronotum, more transverse on elytra. Head:
eyes moderately prominent (abruptly so
in distorta), with 2 seta-bearing pimctures
over each eye except anterior puncture
absent in distorta and reduced in aculeata
group to a small impressed puncture \\'ith-
out seta; clypeus c. truncate, 1-setose each
side; labrum rather long, subtruncate or
slightly arcuate at apex except slightK
emarginate in microdera, 6-setose; mentum
without tooth, at most slighth' arcuatcK
prominent at middle; ligula and maxillae
fused, together broadly rounded, \\\ih usu-
ally 2 principal setae slighth' bc>fore apex
and additional smaller setae at apex ( setae
often difficult to distinguish); palpi rath(>r
short, but apical segments not widened; an-
tennae moderate, pubescent from 5th seg-
ments, sometimes a little pubescence on
apex of 4th. Prolhorax: .setac> at basal
angles present, median-lateral setae present
or absent; base not margined at middle,
apex usualK' with fine marginal line entire;
discal impressions usually present, some-
times almost obsolete. Elytra varying in
form and in pi-esence or absence of apical
spines; striation entire, varying in depth
and in pnnctalion; .'^rd inlcrxals nsnall\ w itli
2 niinutt> punctures without setae placed
irregularly in posterior half of elytral length
(see Notes. b(4ow). Inner wings bill. Legs
The Carabid Beetles of New Guinea • Darlington 125
slender; 4th tarsal segments weakly emar- elytra too. The species of this group differ
ginate; 5th segments with few, weak ac- among themselves in color, elytral micro-
cessory setae; claws with c. 3 to 5 teeth, sculpture, and form of elytral striae. This
Secondary sexual characters: 6 front tarsi group is thus far known only from the
very little dilated, 3 segments with paired eastern half of New Guinea, and the species
squamae at least near apex; S middle tarsi are partly allopatric: divisa and hiion seem
without squamae; c^ middle tibiae not ex- not to occur together, and neither do
cised; S with 1, 9 1 or 2 setae each side castanea and polita.
last ventral segment. Although the New Guinean Dolichoctis
Type species. Dolichoctis .striata Schmidt- are rather diverse, they are less so than
Goebel (below). the Oriental members of the genus. Some
Generic distribution. Numerous from Oriental groups, including Menartis (a
SE. Asia across the islands to New Guinea, group of small convex species ) , are not
a few species reaching New Britain, New represented in New Guinea at all. The
Ireland, the Solomons, and northern general pattern of distribution of the genus
Australia. suggests that 3 or 4 stocks have reached
Notes. The 13 New Guinean species of New Guinea at different times, probably
Dolichoctis can be arranged in 5 groups. D. all from the direction of tropical Asia, and
striata and microdera represent separate that 1 or 2 of the older stocks have radiated
species groups which arc widely distributed moderately on the island,
outside New Guinea and which have prob- The 2 minute impressed punctures, with-
ably reached New Guinea comparatively out setae, on each 3rd elytral interval pos-
recently from the west. D. distorta is unique teriorly are present in most Dolichoctis but
and forms a group of its own; it may be may be absent in distorta (in which these
derived from either of the following species punctures, if present, are lost in the general
groups or from a common ancestor. Six punctation of the intervals) and are dif-
of the remaining New Guinean species form ficult to see and perhaps sometimes absent
what I call the aculeafa group, character- in the polita group. These minute punctures
ized by anterior supraocular seta-bearing are best seen in carefully cleaned speci-
punctures reduced to small impressed mens under diffused light. Wlien I have
points without setae, median-lateral pro- been able to see them clearly in the first 1
notal setae lost, elytra dentate or spined, or 2 specimens of a series, I have credited
and reticulate microsculpture present on that species wath possessing them, without
entire upper surface. Excepting dentata, attempting to clean and examine whole
which is satisfactorily distinct, the species series.
of this group are very similar to each other Although my field notes do not distin-
and difficult to define because of occur- guish most species of this genus, I know
rence of intermediates. They are sympatric that most of them (except rriicrodera) are
— all 6 species occur at Dobodura — and arboreal, living in understory foHage in
do not seem to be differentiating geographi- rain forest. They are usually collected by
cally. This group is represented also out- sweeping or beating. They do not often
side New Guinea. Finally, 4 species form fly to light, which suggests that they may
what I am naming the polita group, which be largely diurnal. However, one species,
is like the aculeata group in form and in distorta, is apparently known only from
spined elytra, but anterior supraocular seta- light-collected specimens. It may be noc-
bearing punctures are present, and reticu- turnal and may occupy a habitat that col-
late microsculpture is absent on head and lectors do not often reach, perhaps tree-
pronotum and in some cases absent on tops in rain forest.
126 BitUetin Museum of Comparative Zoology, Vol 137, No. 1
Key to Species of Dolichoctis of New Guinea
1. EKtia obliquely truncate at apex, not
spined or denticulate; elytra usually
spotted 2
- Elytra spined or acutely denticulate at
apex; not spotted, except sutural area some-
times red 3
2. Prothorax wide and widely margined (p.
126 ) striata
- Prothorax narrow, narrowly margined (p.
127 ) inicrodera
3. Head distorted, eyes small but abniptly
prominent, front swollen on each side;
prothorax semicircular, more than 2X wide
as long ( p. 127 ) distorta
- Head normal; prothorax less than 2x wide
as long _^ 4
4. Head and pronotum (and elytra) with
reticulate microsculpture; anterior seta-
bearing punctures over eyes reduced to
minute punctures without setae {acidcata
group ) 5
- Head and pronotum without reticulate
microsculpture ( elytra with or without it ) ;
anterior ( as well as posterior ) seta-bearing
punctures present over eyes ( polita
group ) 10
5. Form broader, more Agomim-MVe (Fig.
82 ) ; prothorax relatively smaller and nar-
rower; elytra dentate at apex (p. 128) -._.
dentata
- Form more oval or fusiform; prothorax
usually relatively larger and wider; elytra
spined at apex 6
6. Elytral striae very lightly impressed, 7th
striae reduced to very fine superficial
lines; elytral spines usually very long (but
varial)le) (p. 128) .v/j/'/io.sr/
- Elytral striae including 7th well impressed;
elytral spines usually shorter 7
7. Suture or sutural area red; form usually
relatively narrow; length 4.6-5.6 mm. (p.
129) sutundis
- Suture not red; form variable; size often
larger - __ _ 8
8. Sides of prothorax ± strongly sinuate;
elytral striae moderately impressed (p.
129) .._^ andcala
- Sides of prothorax not or only .slightly
simiale; elytral striae often ilceper 9
9. i'rothorax wider (width/length 1.78 and
1.80), with sides more rnuiidcd and wiUi
wider margins (p. 130) siihrottiiKia
- Prothorax narrower (width length 1.61 and
1.69), with sides less rounded and with
narrower niargins (p. 130) sidxpiddrdtd
10. Strikingl)- bicolorcd, head and prothorax
red, elytra black or piceous; often larger
(5.8-7.4 mm) II
- Not or at most \aguely bicolored; often
smaller ( 5.4-6.5 mm ) ._„ 12
11. Elytra with grooved striae and distinct
reticulate microsculpture (p. 131) .._. divi.sci
- Elytra with striae formed by rows of small
punctures and elytral disc without reticu-
late microsculpture (p. 131) huon
12. Elytra with reticulate microsculpture (p.
131 ) castanea
- Elytral disc without reticulate microsculp-
ture (p. 132) polita
Dolichoctis striata Schmidt-Goebel
Schmidt-CIoebel 1846, Faunula Coleop. Birmaniae,
p. 62.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1384 (see for synon\'my and additional refer-
ences ) .
Louwerens 1958, Treubia 24, p. 258.
Jedlicka 1963, Ent. Abhandlungen 28, p. 357.
Description ( for recognition only ) . With
characters of genus; form broad, with elytra
unarmed; black or piceous, elytra usually
with red spots; 2 setae over each eye; pro-
thorax with 2 setae each side (see Notes,
below); length c. 4—4.5 mm.
Ttjpe(s). From Burma, in Prague Mus.
(not seen).
Occurrence in Netv Guinea. Common
probably throughout New Guinea: 200
specimens (by count), including examples
from Normanbv, Woodlark, Biak, and \Vai-
geu Is.; most at low altitudes (including
Dobodura), but a few at 1050, 1100, and
1200 m at Wau.
Notes. The recorded range of striata is
from SE. Asia (including Ceylon and
Japan) to the Philippines, Ne^ (ruinea.
and Xorth Queensland, Australia, ;uid I
have specimens also from New Britain and
New Ireland. \Miether populations from
all these places are in hut conspecific is
a (]uestion for future study.
Most individuals from New CJuinea are
either 4-spottcd (each clvtron with a pale
spot near bas(> and another near apex), 2-
spottcd (with oiiK the subapical spots), or
intermediate (with consiiicuons subapical
and fainter subb;isal spots — note that the
subbasal elytral spots \ ;ny in distinctness
more than in si/(0. The siiiule iiidiv idujil
The Carabid Beetles of New Guinea
Darlington
127
seen from Woodlark Is. is the only un-
spotted one in the New Gninean series.
However, 4-spotted, 2-spotted, and un-
spotted individuals are said to occur else-
where in the species' range (Louwerens
1958).
Of the 200 New Guinean individuals, all
that are in condition to examine have 2
seta-bearing (or formerly seta-bearing)
punctures over each eye, and all have both
a basal and a median-lateral seta (or
puncture) in each prothoracic margin ex-
cept that 2 specimens from Nabire, West
N. G. (Bishop Mus. ), lack the median-
lateral seta and puncture on one side.
These specimens have the pronotum slightly
unsymmetric: angulate on the side with
median seta, evenly arcuate on the side
without seta. Numerous other specimens
from the same locality have the median
seta and puncture present on both sides.
This species lives in understory foliage
in rain forest.
Dolichoctis microdera Andrewes
Andrewes 1930, Ann. Mag. Nat. Hist. (10) 6, p.
665.
1931, Zoologische Mededelingeii 14, p. 63.
Louwerens 1956, Treubia 23, p. 226 ( Moluccas ) .
1964, Ent. Tidskrift 85, p. 184 (Borneo).
Description ( for recognition only ) . With
characters of genus; form (Fig. 80) rela-
tively slender, with narrow, narrowly mar-
gined prothorax; black or piceous, each
elytron with 2 pale spots; 2 setae over each
eye; prothorax with basal but not median-
lateral setae; length c. 4.5-5 mm.
Type. From Sumatra; in Andrewes
Coll., British Mus. (seen).
Occurrence in New Guinea. Papua: 6,
Dobodura, Mar.-July 1944 (Darlington).
N-E. N. G.: 1, Nadzab, July 1944 (Dar-
lington); 1, Torricelli Mts., Siaute, sea
level, Nov. 9-17, 1958 (W. W. Brandt,
Bishop Mus.).
Notes. Comparison (made in 1948)
shows that New Guinean specimens differ
slightly from the Sumatran type, but the
latter is unique. More material from more
localities is needed to show whether the
differences are individual or geographic.
The known range of the species now in-
cludes Sumatra, Borneo, the Moluccas,
and New Guinea.
My New Guinean specimens were (I
think) taken among dead leaves on wet
ground, a unique habitat for members of
this eenus in New Guinea.
Dolichocfis distorta n. sp.
Description. With characters of genus;
form as in Figure 81; irregularly reddish
piceous, appendages irregularly brown;
shining, dorsal reticulate microsculpture
lacking but most of surface irregularly,
rather finely punctate. Head 0.64 and 0.65
width prothorax; eyes abnormally small but
abruptly prominent, with a channel over
each eye running diagonally forward; pos-
terior seta-bearing puncture high above
each eye, anterior puncture absent; front
strongly swollen each side of median longi-
tudinal channel, each swollen area im-
pressed near middle; sides of head behind
eyes longitudinally multisulcate. Frothorax
very wide; width/length 2.13 and 2.18;
base/apex 1.43 and 1.40 (base measured
across seta-bearing punctures); sides very
broadly rounded into base, with posterior
angles not defined; margins very widely
depressed, slightly reflexed; posterior-lateral
setae present, median-lateral setae absent;
base and apex not margined; disc with
median line and posterior and anterior trans-
verse impressions. Elytra: width elytra/
prothorax 1.16 and 1.17; outer-apical angles
distinct, c. right or minutely acute; apices
each with short spine c. opposite ends 2nd
intervals; striae impressed, not punctate;
intervals punctate, 3rd with usual 2 small
punctures doubtfully distinguishable behind
middle. Secondary sexual characters as for
genus; $ with 1, 9 2 setae each side last
ventral segment. Measurements: length
5.7-6.5; width 2.5-2.7 mm.
128 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Types. Holotype 9 (M.C.Z., Type No.
31,424) from Dobodura, Papua, Mar.-July
1944 (Darlington); 3 paratypes, Kokoda,
Papua, Mar. 28-29, 1956 (Gressitt), taken
in light trap; and 1 paratype, same locality,
1200 ft., June 1933 (Cheesman).
Measured specimens. A i paratype from
Kokoda ( British Mus. ) and the 9 holotype.
Notes. In spite of its unique modifica-
tions, this species is clearly a Dulichoctis.
All known specimens were probably taken
at light: Gressitt's are so labeled; Miss
Cheesman's specimen has the scales of
Lepidoptera stuck to it; and mine was
taken on a lighted window.
Dolichocfis acuieafa group
Dolichoctis aculeata Chaudoir and its
immediate relatives form a well defined
group with the following characters in addi-
tion to characters of the genus: form usu-
ally suboval or fusiform ( but broad
AiS,onum-\\Ve in dentata); entire upper sur-
face microreticulate; posterior seta-bearing
punctures over eyes present, anterior pimc-
tures reduced to minute points without
setae; prothoracic margins with seta-bearing
punctures at basal angles, without median-
lateral punctures; elytra with outer-apical
angles well defined (except in dentata)
and apices acutely dentate or spined c.
opposite ends of 1st striae or 2nd intervals;
last ventral segment with 1 seta each side
in both sexes.
Besides aculeata itself (as I identify it),
the lollowing 4 closely related new species
occur in New Ciuinea: sj)in()sa, suturalis,
suhrotiinda^ and subijuadrata. These 5
species (including (/rj//rY//r/ ) apparenlK iii-
tergrade to some extent, and their status
is tliereh)re doubtlul. D. dentata is more
distinct. The species of this group arc all
sympatric in New Guinea.
Although most species ot the acideata
group are New Guinean, the group is repre-
sented west at least to ('elebes (by typical
aculeata), on New Britain, New Ireland,
and the Solomons, and in North Queens-
land, Australia. The group is apparenth'
not represented in the Philippines.
Dolichocfis dentafo n. sp.
Description. With characters of genus
and of aculeata group; form as in Figure
82; broad-Ai;o;Hn?!-like with relati\ely small
prothorax; brownish black, margins and
legs paler brown, antennae and mouthparts
testaceous. Head 0.82 and 0.83 width pro-
thorax. Protliora.x rather small, quadrate-
subcordate; width length 1.58 and 1.60;
base apex 1.23 and 1.21; side margins
moderateK' wide and reflexed. Elytra:
width elytra prothorax 1.71 and 1.73; outer-
apical angles rounded, apices acutely den-
tate; striae moderately impressed, impunc-
tate. Measurements: length 6.5-7.0; width
2.8-3.1 mm.
Types. Holotype 6 (M.C.Z., Type No.
31,425) and 21 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
6 additional paratypes from Oro Bav
(near Dobodura), "Dec. 1943-Jan. 1944
( Darlington).
Additional material. N-E. N. G.: 1, Sur-
prise Ck., Nh)robe Dist., Oct. 7 (Stevens,
M.C.Z.); 1, Simbang, Huon Gulf, 1898
(Biro); 2, Torricelli Alts., Wantipi Village,
Nov. 30-Dec. 8, 1958 (W. W. Brandt,
Bishop Mus.). West N. G.: 1. Ilollandia,
Jan. 1945 (Malkin, U.S.N.M.); 1, llijob, 25
m, Sept. 10, 1956 ( Neth. New Guinea Exp.,
Leiden Mus.); 1, Wasian, \\)gelkop, Sept.
1939 (Wind, M.C.Z.).
Measured specimens. The i holotvpe and
1 9 paratype from Dobodura.
Notes. I know no \c'r\' close relati\es of
this species. It is, ol course, placed in rela-
tion to others in tlu' iireceding Key lo
Species.
Dolichocfis spinosa n. sp.
Description. With characters of genus
and ol (icutcala gioup; black. ai')pendages
dark brown; niicrosculpturc more transverse
than usual on pronotum and elytra, latter
sliuhtK iiidcsceut. Head 0.74 and 0.76
The Carabid Beetles of New Guinea • Darlington
129
width prothorax. Prothurax transverse-cor-
date; width/length 1.67 and 1.64; base/apex
1.26 and 1.25; sides depressed but margins
not well defined. Elytra: width elytra
prothorax 1.61 and 1.72; outer-apical angles
well defined but obtuse, apices with long
(but variable) slightly dehiscent spines;
striae, especially outer ones, very lightly
impressed. Measurements (types only);
length c. 6.0-7.5 (including spines); width
2.6-3.2 mm.
Types. Holotype $ (M.C.Z., Type No.
31,426) and 9 paratypes from Dobodura,
Papua, Mar.-July 1944 (DarHngton). Ad-
ditional paratypes as follows, all from
Papua: 3, Kokoda, 1200 ft., June, Aug.,
Sept. 1933 (Cheesman); 1, Palmer R. at
Black R., June 7-14, 1936 (Archbold Exp.,
A.M.N.H.).
Additional material. Eighteen from vari-
ous localities in all 3 political divisions of
New Guinea; some at low altitudes, some
at 1200 (at \\'au), 1300, and 2000 m.
Measured speeimens. The i holotype and
1 9 paratype from Dobodura.
Notes. Typical specimens of this new
species are easily recognized by very light
elytral striation and very long elytral spines,
but some individuals listed under Additional
material have shorter spines and vary
toward one or another of the following
species.
Dolichoctis sufuralis n. sp.
Deseription. With characters of genus
and of aeuleata group; form c. as in aculeata
(following species) but more slender,
smaller; castaneous with suture or sutural
area reddish, appendages brownish testa-
ceous. Head 0.69 and 0.74 width prothorax.
Prothorax: width length 1.79 and 1.74;
base/apex 1.40 and 1.39; sides broadly
rounded anteriorly, slightly sinuate before
somewhat obtuse, blunted posterior angles;
margins rather widely depressed especially
posteriorly. Elytra rather narrow; width
elytra prothorax 1.44 and 1.52; outer-apical
angles well defined but obtuse, apices
with short spines; striae moderately im-
pressed, not distinctly punctate. Measure-
ments: length c. 4.6-5.6; width e. 2.0-2.4
mm.
Types. Holotype i (M.C.Z., Type No.
31,427) and 23 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
6 paratypes from Oro Bay (near Dobodura),
Dec. 1943-Jan. 1944 (Darlington).
Additional material. Thirty-one (some
doubtfully identified), from numerous lo-
calities including all 3 political divisions of
New Guinea and Normanby Is.; most from
low altitudes ( usually below 500 m ) but 1
from Finisterre Rge. at 1200, and 1, Upper
Jimmi Vy. at 1300 m.
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. In the aculeata group of New
Guinean Dolichoctis, only relatively small,
slender individuals have reddish sutures.
This correlation of size, form, and color
suggests that suturalis is a real species, al-
though the distinguishing characters are
slight.
Dolichoctis aculeata Chaudoir
Chaucloir 1869, Ann. Soc. Ent. Belgium 12, p.
251.
Andrewes 1930, Treubia 7, Supplement, p. 336.
Louwerens 1956, Treubia 23, p. 226 (Moluccas).
Description. With characters of genus
and of aculeata group. Head 0.72 and 0.67
width prothorax. Prothorax: width/length
1.80 and 1.75; base/apex 1.30 and 1.42
( proportions notably variable ) ; sides rather
broadly depressed. Elytra: width elytra/
prothorax 1.50 and 1.49; outer-apical angles
well defined, almost right ( slightly obtuse ) ,
apices spined; striae moderately impressed.
Measurements: length c. 5.0-6.5; width c.
2.2-3.0 mm.
Types. From Celel>es, collected by
W^allace; type now in Oberthiir Coll., Paris
Mus. (not seen).
Occurrence in New Guinea. Common
probably throughout the island: 120 speci-
mens (including 65 from Dobodura and
Oro Bay), from all 3 political divisions of
130 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
New Guinea and Rossel and Woodlark Is.;
most at low altitudes but reaching 1200 to
1400 m at some localities.
Measured specimens. A pair { i 9 ) from
Dobodura.
Notes. My identification of this species
is based on comparison with specimens
identified by Andre wes in his collection.
I collected speciinens that I refer to this
species at Iron Range and Rocky R. in
the mid-peninsular rain forest of Cape
York, Australia, in 1958.
Dolichoctis subrotunda n. sp.
Description. With characters of genus
and of aculeata group; similar to oculeata,
differing principally in form of prothorax
(Fig. 83), with broadly rounded sides not
or scarcely sinuate posteriorly. Head 0.65
and 0.66 width prothorax. Prothorax:
width/length 1.78 and 1.80; base/apex 1.42
and 1.33; sides flattened but not strongly re-
flexed. Elytra: width elytra/prothorax 1.49
and 1.49; apical spines moderate; striae mod-
erately impressed. Measurements (types):
length c. 5.0-6.5; width c. 2.3-2.9 mm.
Types. Holotype 6 (M.C.Z., Type No.
31,428) and 26 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
2 additional paratypes from Oro Bay ( near
Dobodura), Dec. 1943-Jan. 1944 (Dar-
lington ) .
Additional material. Sixty (some doubt-
fully identified) from numerous localities
in all 3 political divisions of New Guinea
and Normanby and Woodlark Is.; most at
low altitudes, but recorded above 1000 m
at several localities including Wau and at
2500 m in the Chimbu area.
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. Individuals assigned to this spe-
cies vary considerably in si/e, depth of
elvtral striae, etc. A single specimen from
Waigeu Is. (Camp Nok, 2500 ft. (c. 770
m), Apr. 1938, Cheesman) differs from all
specimens from the mainland of New
Cuinea in having a poorly defined sub-
apical red spot on (>ach elytron near suture.
Doiichocfis subquadrato n. sp.
Description. With characters of genus
and of aculeata group; form similar to
aculeata except prothorax smaller and sub-
quadrate (Fig. 84). Head 0.75 and 0.70
width prothorax. Prothorax: width length
1.61 and 1.69; base apex 1.36 and 1.44;
sides usually slightly sinuate near base;
margins scarcely depressed anteriorly, more
broadly so posteriorly. Elytra: width elytra/
prothorax 1.60 and 1.54; apices with mod-
erate spines; striae usually deeply impressed
(deeper than in aculeata); 7th intervals
slightly elevated at base. Measurements
(types only): length 5.7-6.7; width 2.5-2.8
mm.
Types. Holotype S (M.C.Z., Type No.
31,429) and 3 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); 2
additional paratypes from Oro Bay (near
Dobodura), Dec. 1943-Jan. 1944 (Darling-
ton); and 4 paratypes from Milne Bay,
Papua, Dec. 1943 (Darlington).
Additional material. Four from widely
scattered localities in New Guinea; and 3,
Aru Is. ( British Mus. ) . Also several doubt-
fully identified from Wau.
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. This species seems clearly distinct
from the preceding one (subrotunda): the
proportions of head prothorax and pro-
thoracic width length reflect the relativeK'
large, wide prothorax of subrotunda and the
smaller and narrower one of subquadrata.
However, aculeata is intermediate. These
3 species together form a bewildering, vari-
able complex that includes many individ-
uals wliieh I cannot ]ilace satislactoriK'.
Dolichoctis polifa group
The following four species lorni a group,
apparently confined to Ni'w Cuinea, char-
acterized as follows: form c. as in aculeata
group l)ul more skMider; microsculpture
absent on head and pronotum, prc\scMit or
absent on elytra; 2 setae over each eye;
pronotum with setae* (or inmetures) at basal
The Carabid Beetles of New Guinea • Darlington 131
angles but median-lateral setae absent;
elytral apices dentate or spined; 1 seta
each side last ventral segment in both sexes.
Dolichoctis divisa n. sp.
Description. With characters of genus
and of polita group; form as in Figure 85;
head and prothorax red, elytra black and
slightly silky, legs dark, antennae pale;
elytra with transverse microsculpture. Head
0.63 and 0.67 width prothorax. Prothorax:
width/length 1.52 and 1.57; base/apex 1.41
and 1.38; sides slightly sinuate before c.
right (narrowly rounded) basal angles;
sides of disc slightly depressed. Elytra:
width elytra prothorax 1.35 and 1.37; outer-
apical angles c. right, apices with moderate
spines; striae well impressed, not punctate.
Measurements: length 6.6-7.4; width 2.6-
2.9 mm.
Types. Holotype $ (M.C.Z., Type No.
31,430) and 1 9 paratype from Dobodura,
Papua, Mar.-July 1944 (Darlington), and
additional paratypes as follows. Papua: 2,
Bisianumu, E. of Port Moresby, 500 m,
Sept. 23, 1955 (Gressitt); 1, Kokoda, 1200
ft. (366 m), Aug. 1933 (Cheesman); 1,
Milne Bay, Dec. 1943 (Darlington); 1,
Brown River, 20 km N. of Port Moresby,
Apr. 29, 1960 (C. W. O'Brien, Bishop
Mus.); 1, Popondetta, 60 m, Oct. 18, 1963
(Shanahan, Bishop Mus.); 4, Mt. Laming-
ton, 1300-1500 ft. (c. 400-460 m) (C. T.
McNamara, S. Austrahan Mus.).
Measured specimens. The pair ( i 9 )
from Dobodura.
Notes. This strikingly bicolored species
is apparently confined to a small part of
eastern New Guinea.
Dolichoctis liuon n. sp.
Description. With characters of genus
and of polita group; head and prothorax
red, elytra usually darker ( castaneous ) ,
sometimes scarcely darker; legs and anten-
nae dark; whole upper surface without
reticulate microsculpture. Head 0.65 and
0.66 width prothorax. Frothorax: width
length 1.66 and 1.67; base/apex 1.40 and
1.41; sides slightly sinuate before slightly
obtuse (nearly right) basal angles; sides of
disc slightly depressed. Elytra: width
elytra /prothorax 1.39 and 1.40; outer-apical
angles c. right, apices short-spined or
acutely toothed; striae scarcely impressed,
formed by rows of small punctures. Mea-
surements: length 5.8-6.9; width 2.3-2.8
mm.
Types. Holotype i (Bishop Mus.) and
6 paratypes from Pindiu, Huon Pen., N-E.
N. G., 500-600, 750-850, 870-1300 m, Apr.
19, 20, 21, 21-22, 1963 (Sedlacek). Addi-
tional paratypes as follows, all from north-
ern part of N-E. N. G.: 4, Finschhafen, 10,
80 m, Apr. 12, 16, 1963 (Sedlacek); 1, Lae,
10 m. May 12, 1966 (Gressitt); 1, Busu R.,
E. of Lae, 100 m, Sept. 15, 1955 (Gressitt);
1, Torricelli Mts., Mobitei, 750 m. Mar. 5-
15, 1959 (W. W. Brandt, Bishop Mus.).
Some paratypes now in M.C.Z. (Type No.
31,431).
Measured specimens. The S holotype and
1 9 paratype from Finschhafen.
Notes. The dark antennae, punctate
elytral striae, and absence of elytral micro-
sculpture clearly distinguish this species
from divisa (above). These 2 species are
apparently allopatric, confined to different
small areas of eastern New Guinea.
Dolichoctis costonea n. sp.
Description. With characters of genus
and of polita group; reddish castaneous,
prothorax sometimes slightly paler, append-
ages reddish brown; elytra with transverse
reticulate microsculpture. Head 0.66 and
0.68 width prothorax. Prothorax: width/
length 1.62 and 1.63; base/apex 1.32 and
1.31; sides broadly rounded, not or slightly
sinuate before usually obtuse basal angles;
disc slightly depressed at sides. Elytra:
width elytra prothorax 1.33 and 1.45; outer-
apical angles distinct but obtuse and slightly
blunted, apices short-spined or acutely
dentate; striae well impressed, not distinctly
punctate. Measurements: length 5.4-6.5;
width 2.1-2.5 mm.
Types. Holotype 6 (M.C.Z., Type No.
132
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
31.432) and 2 paiatypes from Dobodiira,
Papua, Mar.-July 1944 (Darlington); and
additional paratvpes as follows. Papua:
3, Kokoda, 1200 ft., July, Aug. 1933
(Cheesman); 1, Kokoda-Pitoki, 450 m. Mar.
24, 1956 (Gressitt); 2, Mt. Lamington, 1300-
1500 ft. (c. 400-460 m) (C. T. McNamara,
S. Australian Mus.); 1, Brown R., May 25,
1956 (E. J. Ford, Jr., Bishop Mus.). N-E.
N. G.: 15, Pindiu, Huon Pen., 500-600,
860, 870-1300 m, Apr. 19-22, 1963 (Sed-
lacek); 1, Busu R. E. of Lae, 100 m, Sept.
14, 1955 (Gressitt); 1, Bubia, Markham Vy.,
50 m, Sept. 20, 1955 (Gressitt); 1, Sattel-
berg, Huon Gulf, 1899 (Biro); 1, Madang
(Friedrich-Wilh.-hafen), 1901 (Biro).
MecLsurcd specimens. The S holotype and
1 9 paratype from Dobodura.
Notes. This speeies is sympatrie with the
two preceding ones, but perhaps allopatric
with the following (polita).
Dolichoctis polifa n. sp.
Description. With characters of genus
and of polita group; reddish castaneous,
prothorax sometimes slightly paler, append-
ages not or slightly paler; entire upper
surface without reticulate microsculpture.
Head 0.66 and 0.66 width prothorax. Vro-
thorax: width/length 1.64 and 1.78; base/
apex 1.38 and 1.36; sides usually not sinuate
before obtuse, sometimes blunted posterior
angles; sides of disc moderately depressed.
Elytra: width elytra/ prothorax 1.33 and
1.36; outer-apical angles distinct but obtuse
and slightly blunted; apices short-spined or
acutely toothed; striae well impressed, im-
punctate. Measurements: length 5.5-6.5;
width 2.1-2.6 mm.
Tijpes. Holotype i (Bishop Mus.) and
33 paratypes (some in M.G.Z., Tvp(> No.
31.433) from Wan, Morobe Dist, N-E.
N. G., altitudes from 1050 to 1500 m, dates
in Jan., Feb., Mar., June, July, Sept., Oct.,
Dec, 1961-1964 (Scdlacek);' and 2 para-
types, Upp(>r Watut R., 24 km W. oi
Bulolo, N-E. N. G., 760 m. Mar. 5-6, 1963
(Scdlacek).
Measured specimens. The i holotype and
1 9 paratype from Wau.
Notes. This may be a geographic form
(confined to the Morobe area) of Dolicho-
ctis castanea (aboye), distinguished pri-
marily by absence of clytral microsculp-
ture, but I do not care to recognize sub-
species in this genus until relationships and
geographic patterns are better understood.
Genus STRICKLANDIA Macleay
Macleay 1886, Proc. Linnean Soc. New South
Wales (2) 1, p. 138.
Diagnosis. Form ( Fig. 86 ) characteristic,
large, yery broad, depressed; prothorax
strongly cordate, with numerous extra lat-
eral setae anteriorly; elytra yery wide, each
2-spined. See also Key to Genera of Lebiini
of New Guinea.
Description. Form broad, depressed;
black, moderately shining; not obyiously
pubescent but pronotum and sometimes
other parts of upper surface yery incon-
spicuously sparsely setulose; reticulate mi-
crosculpture absent or indistinct on head
and disc of pronotum ( but pronotal disc
transyersely rugulose), yisible but meshes
imperfect and irregular on elytra. Head:
eyes rather small but prominent; 2 setae
oyer each eye; front flattened, weakl\- de-
pressed; clypeus subtruncate \\ ith rounded
angles, 1-setose each side; labrum long,
apex subtruncate or slighth broadly emar-
ginate, 6-setose; mandibles moderately long,
not strongly arcuate, longitudinalb' striate
aboye at middle oF length; antennae slender,
pubescent from middle 4th segments;
mentum subtruncate in sinus, sli^htK' lobed
or with short blunt tooth; ligula wide at
apex, with 2 or 3 large and 2 or more smaller
setae, and paraglossae attached to and
slightly longer than ligula, without setae;
palpi slendei-, apical segments labial palpi
with longitudinal row of numcMous setae
ab()\-e. Prothorax cordate; base not lobed
but irregularly obliciucK rounded to basal
angles; sides angulate or scalloped at mid-
dle. r(41exed, with priiuipal setae at basal
The Carabid Beetles of New Guinea • Darlington 133
angles and near middle and several addi-
tional often smaller setae anteriorly; median
longitudinal and basal and apical transverse
areas impressed; base and apex not dis-
tinctly margined. Elytra very wide, widest
near base; humeri rounded but very promi-
nent; outer-apical and sutural angles both
spined; margins finely serrate and setulose;
striae entire, punctate; 3rd intervals with 1
or 2 seta-bearing punctures behind middle.
Inner winfis full. Lefis slender; 4th seg-
ments middle and hind tarsi narrow, scarcely
emarginate; 5th segments with accessory
setae minute (vestigial?); claws with c. 4
small teeth, in basal half of claw length.
Secondary sexual characters: 6 front tarsi
scarcely dilated but 3 segments with small
2-seriate squamae; S middle tarsi without
squamae; i middle tibiae not excised; S
with 1, ? 2 setae each side last ventral seg-
ment.
Type species. Stricklandia pericalloides
Macleay.
Generic distribution. New Guinea (2 or
more species ) ; Moluccas ( 1 species, from
Batjan Is., Louwerens 1956, Treubia 23, p.
241 ) ; New Britain ( 1 probably unde-
scribed species); and North Queensland,
Australia (1 species). The members of this
genus that I have collected live on tree
trunks and fallen logs in rain forest.
Notes. I do not know the relationships or
geographic origin of this primarily New
Guinean genus.
Key to Species of Stricklandia of New Guinea
1. Prothorax narrower (usually c. 1.5 X wide
as lonfj; at middle, but sometimes wider),
with relatively narrow margins (reflexed
margins often less than V4 as wide as distance
from midline to lateral trough, but some-
times wider) (p. 133) pericalloides
- Prothorax very wide (c. 1.9x wide as long
at middle), with very wide margins (re-
flexed margins more than V2 as wide as
distance from midline to lateral trough)
(p. 133) hta
Stricklandia pericalloides Macleay
Macleay 1886, Proc. Linnean Soc. New South
Wales (2) 1, p. 139.
Description. See generic Diagnosis and
Description. Head 0.79 and 0.78 width
prothorax. Prothorax: width/length 1.48
and 1.56; l^ase/apex 1.18 and 1.15; reflexed
margins relatively narrow. Elytra: width
elytra/ prothorax 1.55 and 1.54. Measure-
ments (Dobodura series): length c. 11.5-
13.5 (including elytral spines); width 4.5-
5.1 mm.
Type. From Fly R., Papua; presumably
in Macleay Mus., Sydney (not seen).
Occurrence in New Guinea. Common
probably throughout New Guinea: 96
specimens (some doubtfully identified, see
following Notes), from all 3 political divi-
sions of the island; most at low altitudes
but reaching c. 1500 to 2000 m at several
localities including Wau.
Measured specimens. A pair { i 9 ) from
Dobodura, Papua.
Notes. Some individuals tentatively as-
signed to pericalloides have prothoracic
margins relatively wide (but not so wide as
the following species) and may be specifi-
cally distinct, but I do not wish to describe
them at present. Mr. Louwerens may refer
these individuals to a species he will prob-
ablv describe from New Britain.
Stricklandia lata n. sp.
Description. With characters of genus;
form as in Figure 86, extraordinarily wide;
color and surface as described for genus,
but elytral microsculpture more transverse
than in pericalloides. Head 0.64 and 0.68
width prothorax. Frothorax wide-cordate;
width length 1.89 and 1.89; base/apex 1.06
and 0.99; margins very wide {c. V2 wide as
distance from inner edge of margin to
middle line), with outer edge irregular.
Elytra: width elytra prothorax 1.29 and
1.36. Measurements: length c. 15-16; width
c. 6.5 mm.
Types. Holotype S (Leiden Mus.) from
Arabu Camp, Wissel Lakes, West N. G.,
1800 m, 1939 (H. Boschma), and additional
paratypes from Wissel Lakes as follows:
Is (M.C.Z., Type No. 31,434), Digitara,
Oct. 1938 (P. J. Eyma); 1 9 , Wagete, Tigi
134 Bulletin Museuin of Comparative Zoology, Vol. 137, No. 1
L., 1700 m, Aug. 17, 1955 (Gressitt); 3,
Enarotadi, 1850, 1850-1900, 1850-2050 m,
dates in July, Aug. 1962 (Sedlacek).
Measured specimens. The c5 holotype and
9 paratype from Wagete.
Notes. Distinguished from pericaUoides
by much wider prothorax and other dif-
ferences of proportion shown l:)y ratios in
the Descriptions.
Genus PEUOCYPAS Schmidt-Goebel
Schmidt-Goebel 1846, Faunula Coleop. Birnianiae,
p. 33.
Jeannel 1949, Coleop. Carabiques de la Rejiion
Malgache, Part 3, p. 991.
Demetrias Csiki 1932 (in part), Coleop. Cat.,
Carabidae, Haipalinae 7, p. 1386 (see for addi-
tional references ) .
Risoplulits Jedlicka 1963 (not Leach), Ent. Abhand-
lungen 28, p. 401.
Diagnosis. In New Guinea, the form
(Fig. 87), small size (under 5 mm), and
long-lobed 4th tarsal segments are diag-
nostic.
Description. None required here. See
detailed description of following new
species.
Type species. P. sutiiraJis Schmidt-Goebel,
of Burma, etc.
Generic distribution. Southern and east-
ern Asia to the Philippines and New
Guinea ( not Australia ) ; Africa, Mada-
gascar.
Notes. Generic distinctions and applica-
tions of generic names have been confused
in the group of genera to which this genus
belongs. In my present use of Teliocijpas
I am following Jeannel, although I do not
like his multiplication of higher categories.
Peliocypas papua n. sp.
Description. Form as in Figure 87;
brown, appendages slightly paler; not
pubescent; rather shining, reticulate micro-
sculpture lightly impressed and irregular, c.
isodiametric on head and elytra, slightly
transverse on disc of pronotum. Head 0.97
and 0.97 width prothorax (measured at
middle); eyes moderately prominent; 2
setae over each eye; front slightly impressed
at sides between eyes and at sides an-
teriorly; frontal suture indicated but not
impressed; clypeus subtruncate, 1 -setose
each side; labrum transverse, subtruncate
with rounded angles, 6-setose; mentum \\'ith
strong triangular tooth; ligula rounded-
subtruncate, apparently 2-setose, with para-
glossae of c. same length, apparently at-
tached, narrowly rounded, without setae.
Prothorax subquadrate, widest at base, with
anterior angles rounded; width (at middle)/
length 1.15 and 1.24; base apex 1.38 and
1.47; base and apex subtruncate (base
slightly sinuate), not margined; side mar-
gins narrow, broader basally and reflexed
and running into deep baso-lateral impres-
sions, each margin with setae at basal angle
and
V.
from apex; disc with usual me
dian line and transverse impressions and
lightlytransverselystrigulo.se. Elytra: width
elytra /prothorax 2.04 and 2.14; humeri
broadly rounded but not much narrowed;
apices obliquely sinuate-truncate, outer
angles rounded, sutural angles blunted;
striae entire but light, not punctate; 3rd
intervals with 2 conspicuous dorsal punc-
tures c. Vi from base and ^4 from apex.
Inner nings full. Legs slender; 4th seg-
ments of middle and hind tarsi with long
lolies; 5th segments with accessory setae;
claws each with 1 long tooth outside and
2 smaller teeth inside middle of length.
Secondary sexual characters: i front tarsi
with squamae (if present) not clearly dif-
ferentiated; last ventral segment with apex
deeply notched in i , entire in 9 ; c^ w ith
1, 9 2 setae each side apex last ventral
seguKMit. Measurements: length c. 4.0-
4.5; width c. 1.8-1.9 mm.
Types. Holotype S (Hungarian National
Mus.) and 5 paratypes (2 in M.C.Z., Tvpe
No. 31,435) all from Madang ("Friedrich-
\\'ilh.-hafen"), N-E. N. G., 1901 (Biro).
Measured specimens. The i holotype and
1 9 paratype.
Notes. This is the (easternmost species of
a genus or group of genera \'er\' w(41
The Carabid Beetles of New Guinea • Darlington 135
represented in the Orient. In Jedlicka's
(1963, pp. 401-402) key to the species of
"R/.wp/n'/us," papiia rnns to conplet 5 but
fits neither species there named, being
narrower-headed than tinicolor JedHcka
and smaller than vimmcri Jedlicka.
New Britain, New Ireland, the Solomons,
Fiji, Samoa, and New Caledonia (speci-
mens seen from all these islands). It lives
in foliage and may (I think) have been
carried eastward into the Pacific by man,
perhaps in thatching material.
Genus CELAENEPHES Schmidt-Goebel Genus SYNTOMUS Hope
Schmidt-Goebel 1846, Faunula Coleop. Binnaniae,
p. 77.
Csiki 1932, Coleop. Cat., Carabidae, Flarpalinae
7, p. 1412 (see for synonymy and additional
references ) .
Jedlicka 1963, Ent. Abhandlungen 28, p. 399.
Diap}osis-. See Key to Genera of Lebiini
of New Guinea and Description of following
species.
Description. None required here.
Type species. Celaenephes parallelus
Schmidt-Goebel ( below ) .
Generic distribution. That of the single
species.
Notes. I do not know the relationships
of this monotypic genus.
Celaenephes parallelus Schmidt-Goebel
Schmidt-Goebel 1846, Faunula Coleop. Birmaniae,
p. 77.
Van Emden 1937, Stettiner Ent. Zeitung 98, p. 35.
Andrewes 1947, Arkiv for Zool. 38A, No. 20, p. 12.
Louwerens 1956, Treubia 23, p. 225 (Moluccas).
See additional references under genus.
Description ( for recognition only ) . Form
as in Figure 88; slender, with elytral apices
simply rounded-truncate; plain black or
piceous; mentum not toothed; claws not
toothed; length c. 6.5-7.5 mm.
Type(s). From Burma; in Prague Mus.
(not seen).
Occurrence in New Guinea. Common
throughout New Guinea and on Normanby
Is.: 206 specimens, most at low altitudes
but a few up to 1550 and 1700 m; found at
Dobodura and Wau.
Notes. This easily recognized carabid
ranges at least from Ceylon, extreme NE.
India (not peninsular India, according to
Andrewes), Burma, etc. to the Philip-
pines and northern Australia, and east to
Hope 1838, Coleop. Manual 2, p. 64.
Jeanne! 1942, Faune de France, Coleop. Carabiques,
Part 2, p. 1075.
Metabletus Schmidt-Goebel 1846, Faunula Coleop.
Birmaniae, p. 38.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1413 (see for additional references, synonymy,
and list of species ) .
Jedlicka 1963, Ent. Abhandlungen 28, p. 420.
Diafinosis. Known among New Guinean
Lebiini by small size, form (Fig. 89),
mentum with emarginate tooth, and tarsal
claws with 2 or 3 minute inconspicuous
oblique teeth.
Description. None required here.
Type species. Of Syntomtis, Carabus
truncatellus Linnaeus, of Europe; of Metab-
letus, M. obscuroguttatus Schmidt-Goebel,
of Burma, etc.
Generic distribution. Temperate and
tropical Eurasia and across the islands
to North Queensland, Australia; North
Ameriea; parts of Africa.
Notes. Only one, widely distributed
species of this genus reaches New Guinea.
Syntomus quadripunctatus (Schmidt-Goebel)
Schmidt-Goebel 1846, Faunula Coleop. Birmaniae,
p. 39 {Metabletus).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1418 (see for synonymy and additional refer-
ences ) .
Description ( for recognition only ) . With
characters of genus; form as in Figure 89;
black; upper surface dull but not pubescent;
elytra with 3rd intervals 2-punctate; length
c. 3.5 mm.
Type(s). From Burma; in Prague Mus.
(not seen).
Occurrence in New Guinea. N-E. N. G.:
4, Wau, Morobe Dist., 1250 m, dates in
Jan., Feb., Sept. 1961, 1962 (Sedlacek); 1,
136 BiiUetin Museum of Comparative ZooIos,ij, Vol. 137, No. 1
Mt. Missim (near Wau), 1050 m, Dec. 27,
1962 (Sedlaceks); 1, Mt. Missim (Stevens,
M.C.Z.); 1, Finschhafen, Apr. 1944 (E. S.
Ross, Cal. Acad.). West N. G.: 1, Eramboe,
80 km ex Merauke, Jan. 29, 1960 (T. C.
Maa, Bishop Mus.).
Notes. The known range of S. quodri-
punctotus is from SE. Asia including
Ceylon, Burma, and Japan across the
Malay Archipelago to the Philippines,
New Guinea, and the NE. corner of Aus-
tralia. Occurrence in AustraUa is based on
a single teneral 9 that I collected N. of
Mareeba, North Queensland, Feb. 1958.
Genus MICROLESTES Schmidt-Goebel
Schmidt-Goebel 1846, Faunula Coleop. Birmaniae,
p. 41.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1420 (see for additional references, synonymy,
and list of species).
Jeanncl 1942, Faune de France, Coleop. Carabiques,
Part 2, p. 1084.
Mateu 1959, Rev. fran^-aise d'Ent. 26, pp. 135 ff.
( species of tropical Asia ) .
1963, Ann. Mus. R. I'Afrique Central, Ser.
in-8°, No. 121, pp. 1-149 ( niono,ii;raph of
African species).
Jedlicka 1963, Ent. Abhandlunsen 28, p. 425.
Diagnosis. Distinguished among New
CHiinean Lebiini by small size, form (note
genae not swollen, prothorax lobed at base ),
mentum without tooth, and tarsal claws
toothed (teeth few and minute in curtatus).
DcscripiUm. None required here.
Tijpc species. Microlestes inconspicuus
Schmidt-Goebel, of Burma, etc.
Generic distribution. Warm-temperate
and tropical Africa and Evirasia and is-
lands to Australia; North America; scat-
tered records elsewhere.
Notes. Two unr(4ated speci(\s occur in
New Ciuinea, one with Oriental and the
other with apparent Australian relation-
.ships.
Kky io Spi:c:n:s ok Microlestes ok Nkw Guinea
1. Hclativcly broad (protlioracic widtli Iciiutli
1.50); elytra with 2 incomplete pale iastiac;
length 3.8 mm (p. 136) ..._ cinctu.s
- Narrower (protlioracic width /]ensj;th 1.18
and 1.28); dull black, not marked; length
not over 3.2 mm (p. 136) curtatus
Microlestes cinctus n. sp.
Description. Form as in Figure 91; black,
in part slightly brownish, elytra with 2 in-
complete transverse fasciae testaceous, ap-
pendages irregularly brownish, bases of
femora and of antennae slightly darker;
rather shining, upper surface with reticulate
microsculpture of meshes isodiametric on
head and pronotum, less regular and slightly
transverse on elytra. Head 0.79 and 0.77
width prothorax. Prothorax \\'ide-subcor-
date; width length 1.50 and 1.55; base
apex 1.12 and 1.14; sides rather narrowly
margined, each margin with setae at base
and c. % from apex; usual discal impres-
sions present but weak. Elytra: width
elytra/prothorax 1.67 and 1.56; striae lightly
impressed, minutely irregular or faintly
punctulate; intervals sparsely minutely punc-
tulate, 3rd with 1 seta-bearing puncture, on
inner edge c. Vi from apex. Legs: claws
each with c. 4 distinct, oblique teeth. Mea-
surements: length 3.6-.3.8; width 1.7 mm.
Ti/pe. Holotype 6 (Bishop Mus.) from
Feramin, N-E. N. G., 1200-1500 m, Mav
11-22, 1959 (W. W. Brandt); 1 S para-
type, Okapa (Okasa), N-E. N. G., July 8,
1965 (Hornabrook), "pine forest, leaf mold."
Notes. Of other species known to me,
this is most like M. atrifasciatus Sloane of
NE. Australia (base of Cape York Pen. to
northern New South Wales), but the color
pattern is different, the ehtra in atrifasciatus
being testaceous with a dark irregular post-
median fascia and subapical and sublateral
dark spots.
Microlestes curtatus n. sp.
Description. i'"onn as in Figure 90;
slender, \\ itli el\ tra niuch shoiter than abdo-
men; dull brow nish black, apjicMidages dark;
entire upinr surface \\ith reticulate micro-
sciilplure irregular (partK' longitudinal) on
head, slightly transverse on pronotiun and
(4ylra. Head 0.92 and 0.89 w idlh i-)rothorax.
Prothorax narrow-subcordatc; width length
The Carabid Beetles of New Guinea • Darlington 137
1.18 and 1.28; base/apex 1.08 and 1.04;
side margins very narrow, each with setae
at basal angle and c. V4 from apex; disc
with median line impressed, transverse im-
pressions scarcely indicated. Elytra very
short, narrowed anteriorly; width elytra/
prothorax 1.68 and 1.64; striae lightly in-
dicated, sometimes scarcely visible, irregu-
lar but not distinctly punctate; 3rd intervals
with 2 punctures, before middle and c. V-t
from apex. Legs: claws each with c. 2
small oblique teeth, easily overlooked. Mea-
surements: length to apex elytra 2.4-2.6,
to apex abdomen 2.8-3.2; width 1.0-1.1 mm.
Types. Holotype S (M.C.Z., Type No.
31,436) and 28 paratypes all from central
plains of Luzon, Philippine Is., Feb.-Sept.
1945 (Darlington).
Occurrence in New Guinea. West N. G. :
2, Dor(e)y (probably collected by Wallace,
British Mus.; this locality is, of course,
somewhat doubtful ) .
Measured specimens. The S holotype and
1 9 paratype from central plains of Luzon.
Notes. I have based this new species on
Philippine individuals because of doubt
about Wallace's locality "Dorey" (see Part
1 of mv work on New Guinean Carabidae,
pp. 330-331).
M. curtatus is similar to exilis Schmidt-
Goebel but has shorter elytra. This species
(curtatus) with very short elytra is not
represented in the Andrewes Collection and
was evidently not known to Mateu (1959)
or Jedlicka ( 1963 ) . It is unknown in Aus-
tralia.
Genus APRISTUS Chaudoir
Chaudoir 1846, Enumeration des Carabiques . . .
Caucase . . ., p. 42.
Csiki 1932, Coleop. Cat., Carabidae, Haipalinae 7,
p. 1432 ( see for additional references, synonymy,
and list of species ) .
Jeannel 1942, Faune de France, Coleop. Carabiques,
Part 2, p. 1083.
Jedlicka 1963, Ent. Abhandlungen 28, p. 427.
Diagnosis. Very small Lebiini, recogniz-
able (in New Guinea) by form (Fig. 92);
surface not pubescent but all or part (at
least elytra) dull and heavily microreticu-
late; genae not swollen; mentum with
entire tooth; claws not toothed.
Description. None required here.
Type species. Apristus suhaeneus Chau-
doir, of the Caucasus and Mediterranean
region.
Generic distribution. \\^arm-temperate
and tropical Eurasia and the Malay Archi-
pelago to the Philippines and New
Guinea (not Australia); part of Africa
(not Madagascar); North and Central
America, Cuba.
Notes. American species of this genus,
which are the only ones I have collected,
live on the ground, usvially on sand or
gravel near water.
Key to Species of Apristus of New Guinea
1. Color brownish bronze; entire upper surface
dull; length 3.0-3.5 mm (p. 137) ^ hiroi
- Color bluish black; front of head and middle
of pronotimi relatively shining, elytra dull;
lengtli 3.5-3.9 mm (p. 137) sedlaccki
Aprisfus biroi n. sp.
Description. With characters of genus;
form as in Figure 92; brownish bronze,
including appendages; entire upper surface
dull, heavily microreticulate. Head 0.90 and
0.90 width prothorax. Prothorax: width/
length 1.28 and 1.30; base/apex 0.91 and
0.90. Elytra: width elytra/prothorax 1.64
and 1.70. Measurements: length 3.0-3.5;
width 1.2-1.5 mm.
Types. Holotype S (Hungarian National
Mus.) and 5 paratypes (2 in M.C.Z., Type
No. 31,437) all from Madang ("Friedrich-
Wilh.-hafen"), N-E. N. G., 1901 (Biro).
Measured specimens. The $ holotype and
1 9 paratype.
Notes. Similar to A. louwerensi Andrewes
of Java, but with elytra more narrowed
anteriorly and with fainter striae.
Apristus sedlaceki n. sp.
Description. With characters of genus;
form c. as in preceding (biroi) except
sides of prothorax more rounded anteriorly
and much more strongly sinuate c. % from
138 Bulletin Museum of Comparative Zoology. Vol. 137, No. 1
base; l)liiish black, appendages dark; front
of head shining with reticulate microsculp-
tiire faint and fragmentary, middle of
pronotal disc ± shining, rest of upper sur-
face including elytra (except edges of
suture) microreticulate and dull. Head 0.92
and 0.91 width prothorax. ProtJiorax:
width length 1.22 and 1.25; base/apex 0.94
and 0.91. Elytra: width elytra prothorax
1.82 and 2.00. Measurements: length 3.5-
3.9; width 1.5-1.8 mm.
Tifpcs. Holotvq^e 6 (Bishop Mus.) and
2 paratypes ( 1 in M.C.Z., Type No. 31,438)
from Tobo-Salembeng, Huon Pen., N-E.
N. G., Apr. 26, 1963 (Sedlacek); 1 para-
type, Golden Pines, Bulolo, N-E. N. G.,
600 m, Feb. 19, 1962 (Sedlacek); and 1
paratvpe, Zengaren, N-E. N. G., 1500 m,
Apr. 28, 1963 (Sedlacek).
Measured speeimens. The S holotype and
1 9 paratype from Tobo-Salembeng.
Notes. This may be related to A.
cuprascens Bates (described from Japan
and identified from the Philippines by An-
drewes), but the color of sedlaeeki is
bluish rather than cupreous, the front is
more shining than in euprascens, and com-
parison of specimens shows slight differ-
ences of form not worth describing in de-
tail here.
(Genus PLOCHIONUS Latreille & Dejean)
Latrcille & Dcjeaii 1824, llistoire KaturcIU- et
Icono^iaphie Coleop. d'Europe 1, p. 150.
Csiki 1932, Coleop. Cat., Carabidac, Ilarpalinae 7,
p. 1451 (see for additional references, synonymy,
sul)fienera, and list of species).
Jeannel 1942, Faune de France, Coleop. Caralii(iu(S,
Fart 2, p. 10:33.
Diaij,ru)sis. See Key to Genera of Lehii)u.
and Description of following species.
Descriplion. None rccjuired h('rc\
Type species. Caral)u.s pcdlens l^'abricius
( below ) .
Generic distribution. Native in tropical
and subtropical Aiiirrica, with th(> follow-
ing species now c. cosmoiMjIilaii.
Notes. A supposed ciKleiiiic PlocJiiinius
in New Caledonia needs confirmation.
{Piochionus pollens (Fabricius))
Faliricius 1775, Systenia Ent., p. 244 (Carahns).
Britton 1948, Proc. Hawaiian Ent. Soc. 13, p. 237
( Hawaii).
Jedlicka 1963, Ent. Alihandlungen 28, p. 450.
See also references under trenus.
Description ( for recognition only ). Fonu
as in Figm-e 93; brown; not pubescent; $
front and middle tarsi slighth' dilated, 2-
seriately squamulose; S middle tibiae
arcuate, lower edges broadly shallowly
emarginate below near middle of length;
length c. 7-9.5 mm.
Type. From Europe ("Habitat Dres-
dae"), now presumed lost (not seen).
Occurrence in Neic Guinea. Not yet
found, but likely to occiu".
Notes. This species, probably originally
from America, has been carried b\' luan
to most of the warmer parts of the world.
In the Asiatic-Pacific region it is known
from SE. Asia, Sumatra, Java, New Brit-
ain, New Ireland, New Hebrides, Fiji, and
Polynesia including Hawaii.
Genus PARENA Motschulsky
Motschulsky 1859, etude Eut. 8, p. 31.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1453 ( see for additional references, synonymy,
and list of species ) .
Jeannel 1949, Coleop. Carahiciues de la Rejj;ion
Malyache, Part 3, pp. 948, 971.
Jedlicka 1963, Ent. AMiandlunsien 28, p. 439.
Diaii,nosis. See Key to Genera of Lehiini
of New Guinea and Figm-e 94; note form
stout, sm-face not pubescent, 4th tarsal seg-
mcMits long-lobed; length c. 8-10 mm.
Dcscri))lio)i. None required here.
Ty))c six'cies. Paroui hicolor Motschulsky,
ol Java.
Generic dislrihution. Most species in
area from SK. Asia (including Japan) to
northern Australia, fewer in Africa and
Madagascar.
Notes. Tile ') species that ha\-e been
found in New (Guinea represent 3 indepen-
dent, w idel\ distributed stocks, f'urther
stn(l\- is needed to clarify their geographic
Nariation and noinciulatui-e; in\' ]')r(\s(>nt
The Carabid Beetles of New Guinea • Darlington 139
material is inadequate. I shall therefore
treat the species only briefly.
All members of this genus that I know
are winged and arboreal.
Key to Species of Parena of New Guinea
1. Color entirely testaceous (with sometimes
vague posterior elytral cloud pale brown);
tarsi and antennae contrastingly black (p.
139 ) testacea
- Color partly or wholly darker; tarsi and
antennae reddish testaceous 2
2. Color testaceous or reddish testaceous with
very liroad, well defined black elytral fascia
(p.' 139) fasciata
- Color irregular rufo-piceous, without well
defined elytral marking (p. 139) picea
Parena fesfacea (Chaudoir)
Chaudoir 1872, Ann. Soc. Ent. Belgium 15, p. 178.
(Crossoglossa).
Description. None required here; see
preceding Key, length (in New Guinea) c.
10 mm.
Types. From the Deccan, India; now in
Oberthiir Coll., Paris Mus. (not seen).
Occurrence in New Guinea. N-E. N. G.:
4, Wau, Morobe Dist., 1200 m, June 25,
Oct. 11-18, Nov. 19, Dec. 5-6, 1961
( Sedlaceks ) .
Notes. This species is now known from
India, (China?), Sumatra, Java ("variety"
cruralis Andrewes), and New Guinea (not
Australia ) .
Parena fasciafa (Chaudoir)
Chaudoir 1872, Ann. Soc. Ent. Belgium 15, p. 179
( Crossoglossa ) .
Jedlicka 1963, Ent. Abhandlungen 28, pp. 440,
443, fig. 154.
sloanei Csiki 1932, Coleop. Cat., Carabidae, Har-
palinae 7, p. 1455 (new synonymy).
plagiata Macleay 1876, Proc. Linnean Soc. New
South Wales 1, p. 167 (Phlocodromius) (new
synonymy ) .
Description ( for recognition only ) . Form
as in Figure 94; yellow or reddish yellow
with conspicuous, broad, transverse, black
elytral fascia; length (in New Guinea) c.
8-9 mm.
Types. Of fasciata, from the Moluecas,
now in Oberthiir Coll., Paris Mus.; of
plagiata, from Yule Is., Hall Sound, Papua,
in Macleay Mus., Sydney; of sloanei (new
name), as for plagiata Macleay (none seen).
Occurrence in New Guinea. Papua: Yule
Is. (type of plagiata). N-E. N. G.: 1, Lae,
July 1944 (F. E. Skinner, Purdue U. Coll.,
Bishop Mus.); 1, Busu R. E. of Lae, 100 m,
Sept. 13, 1955 (Gressitt); 1, Bulolo, 732 m,
Aug. 18, 1956 (E. J. Ford, Jr., Bishop Mus.),
in light trap; 1, Finschhafen, Huon Pen.,
180 m, Apr. 16, 1963 (Sedlacek); 1.
Mumeng, 600 m, Mar. 10, 1962 (Sedlacek).
West N. G.: 2, Hollandia, 250 ft.. May 4,
Nov. 3, 1944 (Hoogstraal, M.C.Z.).
Notes. I have seen specimens that I refer
to this species from Java, Borneo, the
Philippines (including Luzon), Celebes,
the Moluccas, New Britain, and northern
Australia, as well as New Guinea.
Parena picea (Macleay)
Macleay 1871, Trans. Ent. Soc. New South Wales
2, p. 86 (Phloeodromius).
Description. None required here; see
preceding Key; length (in New Guinea) c.
9-10 mm^.
Types. One specimen from Gayndah,
South Queensland, Australia (probably
now in Macleay Mus., Sydney) is presum-
ably the actual type (not seen), although
Macleay mentions also "a few specimens
from other portions of Queensland."
Occurrence in New Guinea. N-E. N. G.:
1, Wau, Morobe Dist., 1200 m, Feb. 25,
1963 (Sedlaceks). West N. G.: 1, Nabire,
S. Geelvink Bay, 10-40 m, Sept. 1-4, 1962
( Sedlacek ) .
Notes. I have no specimens from Aus-
tralia and have identified the New Guinean
ones from the original description. I also
tentatively assign to this species single indi-
viduals from New Britain and Manus Is.
( Bishop Mus. ) .
Genus ANCHISTA Nietner
Nietner 1856, J. Asiatic Soc. Bengal 6, p. 523.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1455 (see for additional references, synonymy,
and list of species ) .
Jedlicka 1963, Ent. Abhandlungen 28, p. 449.
140 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Diag,nosis. See Key to Genera of Lebiini
of New Guinea, and under following
species.
Description. None required here.
Type species. Lehia hrunnea Wiede-
mann, of India and Ceylon.
Generic distribution. Tlie few known
species are confined to SE. Asia including
Ceylon and Japan, except that one (below)
is widely distributed on the Malay Archi-
pelago and islands of the western Pacific.
Notes. I know nothing about the habitat
or habits of members of this genus.
Anchisfa binofata (Dejean)
Dejean 1(S25, Species Ceneiai Coleop. 1, p. 252
(Plucluoiiiis).
See also references under genus.
Description { for recognition only ) . Form
as in Figure 95; brownish piceous, each
elytron with longitudinal testaceous area
centered before middle; surface not pubes-
cent; 5th intervals with conspicuous seta-
bearing piuicture at base; length c. 8-9 mm.
Type(s). From the Marianas; now in
Oberthiir Coll., Paris Mus. (not seen).
Occurrence in New Guinea. Papua: 1,
Hagita, near Milne Bay, Aug. 10, 1919
(J. T. Zimmer, Chicago Mus.).
Notes. This species has now been foimd
in SE. Asia (India to Japan), the Anda-
man Is., Sumatra, Java, Borneo, the
Philippines, Buru, New Guinea, and the
Marianas. It has probably been dispersed
partly by man.
Genus ENDYNOMENA Chaudoir
Cliaudoir J 872, Ann. Soc. Knt. Bcliiium 15, p. 1S6.
Csiki 1932, Coleop. Cat., Carabidae, llarpalinae 7,
p. 1457 (see for additional references, synonymy,
and list of species ) .
jedlicka 196.3, Ent. Ahliandlungcn 28, p. 308.
Diuii^nosis. See Kcij to Genera of Jj'hiini
of New Guinea.
Descri))lion. None required here.
Type species. Plochionus })radieri Fair-
maire (below).
Generic distribution. SE. Asia including
Japan, with the following species very
widely spread over the islands of the
Pacific presumably carried by man.
Notes. The habitat and habits of this
genus too are unknown to me.
Endynomena pradieri (Fairmaire)
Fairmaire 1849, Revue and Magazine Zool. 1, pp.
34, 281.
See also references under genus.
Description ( for recognition only ) . Form
as in Figure 96; brown or piceous; surface
with short pubescence; length c. 8 mm.
Type. From Tahiti; in Oberthiir Coll.,
Paris Mus. (not seen).
Occurrence in Neiv Guinea. N-E. N. G. :
1, Sepik, Maprik area, 160 m, Aug. 29, 1957
(Hardy, Bishop Mus.), at light.
Notes. This insect has been recorded
from parts of SE. Asia, Sumatra, the Phil-
ippines, Fiji, Samoa, Tonga, New Cale-
donia, Tahiti, Hawaii, and other remote
Pacific islands, and I have seen a speci-
men from New Britain (Bishop Mus.).
Genus DEMETRIDA White
White 1846, Voyage Erebus & Tenor, p. 2.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1459 (as subgenus of Xanthophoca) (see
for additional references).
Britton 1941, Proc. R. Ent. Soc. London (B) 10.
p. 188.
Xanthophoca Chaudoir 1848, Bull. Soc. Nat.
Moscow 21, Part 1, p. 73.
Diaiinosis. Among New Guinean Lebiini
of the same general form (Figs. 97-109)
and si/e (5..5-12.0 mm), the species of
Demetrida are distinguished by tarsi pubes-
cent (si^arsely pilose) above, with 4th seg-
ment loug-lobed and tarsal claws with sev-
eral or many long t{>eth; ligula and
paraglossae joiiucl, rounded-truncate, usu-
ally 4-setose ( sometimes with 2 additional
smaller setae); palpi not widely e.xpand(xl;
and ' middle tibiac> usualK' (not al\\a\s)
with inner edge in apical '-t or '2 ol length
with a row of several low tubercles.
Descriplion ( aiiplieablc to all New
Guinean species). l'\)rni usualK slender
(broadest in itnilalrix) , convex; color di-
verse, brown ()!■ black or metallic', nniloim
The Carabid Beetles of New Guinea • Darlinaton 141
or bicolored or tesselated (but pattern not sharply defined, usually subpunctate, but
geometric and not simply 2-maculate ) ; disc otherwise c. smooth or at most sparsely
upper surface with short or long pubescence punctulate. Elytra with humeri rounded,
or not pubescent; reticulate microsculpture margined; outer apical angles rounded or
variable, rarely present on whole upper angulate or denticulate; actual apices
surface, often present only on elytra, some- obliquely truncate or sinuate-emarginate or
times absent; elytral reticulations c. iso- angulate, denticulate, or spined c. opposite
diametric or slightly transverse when not ends of 2nd intervals or 2nd striae; striae
otherwise described. Head narrower than entire, usually well impressed, sometimes
or wider than prothorax; eyes prominent punctulate; intervals flat or moderately
but varying from species to species; genae convex, 3rd with 1 (in tenuis only) or 2 or
usually shorter than eyes and oblique, more dorsal, usually seta-bearing punctures,
rarely angulately prominent; 2 setae over and 5th intervals rarely with similar punc-
each eye, the posterior often distant from tures; when 3rd intervals 2-punctate, the
posterior comers of eyes; front impressed punctures often Vi or less from base on outer
each side anteriorly, often also flattened edge and V-i or less from apex on inner edge
and/or weakly impressed or subpunctate of intervals, but positions vary; additional
at middle; mandibles short, strongly curved; punctures (if present) on 3rd intervals
clypeus 1-setose each side; labrum trans- sometimes smaller and more irregular than
verse, 6-setose anteriorly, with additional the 2 primary punctures. Inner wings full
smaller depressed setae at rounded angles; in all New Guinean species (reduced in
antennae slender, pubescent from ( part of ) some New Zealand and Australian ones ) .
4th segments, first 3 segments more sparsely Lower surface variable, often sparsely or
or not pubescent; mentum with entire partly pubescent (least so in imitatrix);
tooth; ligula and paraglossae equal in prosternal process variable in profile. Legs
length, united, forming a rounded-truncate slender; tarsi pubescent (sparsely pilose)
structure with 4 principal setae and some- above; 4th tarsal segments very deeply
times 2 additional smaller setae; palpi emarginate, with long lobes; 5th segments
pubescent, not widely expanded. Protliorax with accessory setae; claws each with 3
cordate or quadrate or trapezoidal, often to 8 long teeth ( not counting apex of claw )
(not always) as long or longer than wide, and sometimes additional smaller teeth, the
sometimes (not usually) wider at base number varying from species to species
than at middle (but width of prothorax and also varying a little individually, some-
always measured at widest midpoint in com- times different on the 2 claws of one tarsus,
puting proportions); side margins varying Secondary sexual characters: S front tarsi
in width in different species, each margin not or not much dilated but with 3 segments
with a seta at or before middle and some- narrowly 2-seriately squamulose below ( the
times also a seta at or near basal angle, squamae sometimes disarranged and not
and in seticoUis with additional setae an- obviously 2-seriate ) ; 6 middle tarsi without
teriorly (setae often broken off, but their sexual squamules; either i middle tibiae
positions marked by characteristic punc- each with a row of 3 to 9 low tubercles on
tures ) ; disc convex, more so in some species inner edge in outer % or V2 of length, the
than in others; anterior and posterior mar- tibial edge being thus subsinuate or sub-
ginal lines absent or incomplete, middle serrate in profile ( Fig. 160 ) ( this condition
line very coarse and deep (except finer in called tuberculate-serrate), or S middle
kokoda), but subbasal and subapical trans- tibiae modified in some other way, or S
verse impressions weak or obsolete; baso- middle tibiae either straight or slightly
lateral impressions usually present but not bent-in at apex but without tubercles (Fig.
142 BuUetin Muscuui of Comparative Zoology, Vol. 137, No. 1
161) (see Notes below); last ventral seg-
ment with 2 to 4 apical setae each side in 6
(possibly only 1 each side in 6 tenuis), 3
to 8 or more in 9 ( except only 2 each side
in 9 tenuis), the number in each sex usually
fairly constant in a species but varying
somewhat individually and sometimes un-
symmetric with ( for example ) 2 setae on
one side and 3 on the other in a c5 , or 5 on
one side and 6 on the other in a 9 .
Type species. So far as I know, type
species have not been strictly designated
for either Demctrida or Xanthophoea.
These genera were based on New Zealand
and Australian forms respectively, and
type species should be selected during
work on the New Zealand and Australian
members of the group. I therefore make
no designations now.
Generic distribution. Numerous in Aus-
tralia and New Guinea, fewer in New
Zealand, New Caledonia, New Britain,
and the Moluccas (Amboina, Batjan) (oc-
currence in New Britain and Moluccas
based on undescribed material before me ) .
Notes. As Britton points out, Demetrida
has priority over Xantho))]i(>ea. The genus
as a whole is diverse. Perhaps it can be
usefully divided, but this will require re-
vision of the many Australian species, which
seem to bridge the gap between the flight-
less (ground-living?) Demetrida of New
Zealand and the winged (arboreal) species
of New Guinea.
The New Guinean species of Demetrida
may all be interrelated but different ones
differ remarkably in many details. Varia-
tion of some characters within the genus
is indicated in the preceding Description,
and some species groupings are suggested
in the Key to Sjjecies. llowevt^r, diff entices
in the 6 middle tibiae, which may distin-
guish natural species groups, are worth
describing in more detail. Of the 56 species
of the genus now recogni/ed Ironi New
Guinea, l)oth sexes are known ol 47, onK
the 6 of 4, and onlx' the 9 of 5. In most
species of whieli the ■ is known, and also
in at least some Australian and New Zea-
land species, the i middle tibiae have the
inner edge tuberculate-serrate ( see Descrip-
tion, above). The nimiber of tubercles
varies from c. 3 to 9 in different species
(with some individual variation too), and
the tubercles vary in prominence, being
sometimes poorly developed and difficult
to see. The tuberculate-serrate 6 middle
tibiae probably characterize most Demetrida
throughout the genus' range. However,
variations from this pattern occur among
New Guinean species. In D. ni<j,ripennis
( and perhaps also in prima, of which the i
is unknown) the c^ middle tibiae have the
inner edge weakly 2-emarginate. In i
imitatrix each middle tibia has a long
tubercle on inner edge separated from the
apex by an emargination. And the following
16 species have the i middle tibiae straight
or slightly bent-in ( slightly bent-out in
reversa) at apex but not or not distinctly
tuberculate-serrate: tripuncta, genicula, an-
gulata, reversa, kiunga, recta, rex, brunnea,
fumipes, ni<^ricej)s, saidor, divisa, liumcralis.
viridibasis, mafulu, and sibil.
Because of the large number of species
and because many characters are shared by
related species or convergent in unrelated
ones, most New Guinean Demctrida can be
defined only by combinations of characters.
However, D. imitatrix is unique in fonn
( relatively broad ) and in form of 6 middle
tibiae. D. viti^il is uni(iue in abrupt promi-
nence of eyes. D. kokoda is unifiue in form
and in fineness ol impressed niiddli' line of
pronotum. D. seticoUis is unique (among
the nonpubescent species) in possessing
extra lateral pronotal setae* anteriorly. And
D. tenuis is uni{|ue in sculpture of front, in
having only 1 seta-bearing pvmelure on 3rd
elytral inter\al, and in ha\ing oiiK 1 apical
seta on each side in the •" , and only 2 in
the 9 . Besides these* single species with
iiiiiciue eharaeters, the lollowing pairs or
small groups of species share special char-
acters. Among New Ciuincnm Demetrida,
oiilx rcltild. riridilxisis. and niafiiln have
The Carabid Beetles of New Guinea • Darlingion 143
almost the whole upper surface micro-
reticulate; only tripuncta and genicuJa have
the genae stronghj angulately prominent,
although tenuis and some other species
ha\e the genae subprominent; only scriato
and nubicola (of nonpubescent species)
have special seta-bearing punctures on 5th
elytral intervals; and only nig,ripennis and
perhaps prima have the i middle tibiae
2-sinuate on inner edge.
Some New Guinean species of Dcmetrida
are remarkably variable. Great individual
variation is indicated by differences in
proportions of the Measured specimens of
some species. And Mendelian dimorphism
is suggested in some cases. For example,
dimorphism or polymorphism of color ap-
parently occurs in Demetrida diversa (mark-
ings black or green, legs dark or pale) and
in mafnhi (markings present or absent),
and color differences among some other
species may be Mendelian, and presence or
absence of certain prothoracic and elytral
setae may be Mendelian too, as is the case
among some other Carabidae. (Genetic
dimorphism of these and other Carabidae
will be considered in more detail in Part
IV of my work on the carabid beetles of
New Guinea.) This situation suggests that
the explosive evolution of Demetrida in
New Guinea, discussed below, is correlated
with great genetic variability of some
species, as would be expected. Different
species may still share homologous genes,
and characters that have become stabilized
in some species may still be dimorphic or
polymorphic in other species.
I think that Demetrida is in fact in the
very midst of an evolutionary explosion in
New Guinea. This is suggested by the
diversity of superficial differences among
many apparently closely interrelated species
and by the great variability of some species.
Apparently one or more ancestors have
recently invaded an open or incompletely
occupied habitat in New Guinea — primarily
the low foliage of rain forest — where other
predaceous beetles of this size are few.
This habitat is occupied in other tropical
regions by Carabidae of the genus Calleida,
which many of the New Guinean Demetrida
resemble in size, form, and even color, al-
though the two genera are well differen-
tiated taxonomically by differences in
mouthparts, tarsal pubescence, etc. That
Demetrida and Calleida are geographically
complementary is true but an oversimplifi-
cation. The situation is complicated in
many ways, for example by the presence
of many species of Lebia in some other
tropical regions but few in New Guinea.
The ecology of Demetrida seems con-
sistent with a recent independent radiation
of the New Guinean species. While most
New Guinean species apparently live in
fohage in rain forest, most Australian
species live on shaggy-barked tree trunks
(especially of Eucalyptus trees) in rela-
tively open woodland, and the Australian
tree-trunk forms and the New Guinean
rain-forest-foliage forms have evidently
radiated independently. Although this is
true, it is another oversimplification. A few
northern Australian species of Demetrida
do inhabit rain-forest foliage, but they are
very few, uncommon, and probably ecolog-
ically unimportant. Perhaps they represent
the ancestral stock(s) from which the New
Guinean rain-forest forms have evolved.^
^ The following key characterizes 3 species of
Demetrida from North Queensland, Australia, that
are or may be members of the New Guinean
radiation of the genus. D. angulata, described in
the present paper, is the only species known to be
common to Australia and New Guinea. The other
2 Australian species named in the key are distin-
guished by 1-punctate 3rd intervals from all known
New Guinean si^ecies except tenuis.
Key to Certain Australian Demetrida
1. Elytral apices sinuate-truncate; 3rd intervals
1-punctate 2
- Elytral apices obtusely angulate; 3rd intervals
2-punctate (1 9 , Rocky R. on Cape York;
New Guinea) angulata (n. sp. )
2. Prothorax with posterior-lateral setae (11,
middle Cape York, Cairns, Kuranda, etc.,
vie. Brisbane, Clarence R. )
longicollis Macleay
144 BiiUetin Museum of Comparative Zoology, Vol. 137, No. 1
Also, a few Australian Demetrida live in
long grass, and a few unrelated New
Guinean species (perhaps pallen.s?) may
have invaded grassland independently, at
high altitudes — but this is a guess, based
on the insects' appearance; the actual habi-
tats of the New Guinean species in question
are not recorded. However, these excep-
tions and doubts do not change the general
fact: Demetrida has radiated ± indepen-
dently in the New Guinean rain forest, and
the radiation may be continuing explosively
now. The radiation of these beetles parallels
in some ways the radiation of birds of
paradise in the same forests.
The geographic distribution of different
species of Demetrida in New Guinea is
not yet very well known, and the ecologic
distribution of the species within the rain-
forest-foliage habitat is hardly known at
all. Some species of the genus are ap-
parently localized in parts of New Guinea,
and geographic replacement may occur in
some cases. But many other species are
evidently wide-ranging on the island, and
many species sometimes occur together at
one locality or in a very limited area. For
example, I found 8 species at Dobodura,
at relatively low altitudes. And 19 species
have been found at or near Wau on the
Morobe Plateau, at mid-altitudes. The
genus as a whole ranges in New Guinea
from sea level to or above timber line but
is evidently best represented at mid-alti-
tudes, where it is a]:)parently dominant,
and where most of the strikingly colored
species occur. Some species are evidently
confined to or specialK characteristic of
either low, middle, or high altitudes, and
related species may replace each other at
different altitudes in some cases.
- Prothorax without posterior-lateral setae 3
3. Protliorax narrower ( width length 0.84 and
0.93); front snhearinate (New (liniiea
only) (/('/II//.V n. sp. )
- Prothorax wider (width/length 1.07); front
not snhearinate (1 9, Cairns)
_ ferruginca Cha ndoi r
As to their ecolog\ , the bright color of
some New Guinean Demetrida and the
fewness of individuals taken at light sug-
gest that most species are diurnal. How the
various species that occur together, for ex-
ample at Wau, di\ide the niches within
the rain-forest-toliage habitat can only be
guessed at now. A few may have become
nocturnal. Some species certainly live in
understory vegetation in the rain forest,
but some may live at mid-levels and some
77uiy live in the actual tree tops. Different
species tmiy specialize in narrower habitats,
or they 7nay specialize in different kinds
or sizes of prey. But I should repeat that
this is mainly guesswork. There is an op-
portunity here for exciting work in the
field, on the ecologic radiation of a donii-
nant group of insects that is radiating
structurally.
Some New Guinean Demetrida may be
mimics. Evolution of mimetic relationships
would, I think, be consistent with the genus
being now in the midst of an evolutionary
explosion, with many species geneticalh
variable, read)' to respond to special selec-
tion pressures. Demetrida imitatri.x re-
sembles and may mimic the common New
Guinean carabid Viola <i,onum violaeeum
(Ghaudoir), and some other brighth- colored
Demetrida ma\' mimic other (^arabidae
(perhaps certain C^olliurini) and other
Ix'etles.
An extraordinar\ eircumstanee is that,
although many species of Demetrida occur
in New Guinea and although some of them
are common ( 1 have examined a total of
about 1250 individuals) all 56 New Ciuinean
si)ecies seem to be undeseribed! Howe\er,
this should not be interpretc^d as e\id(Mice
ol e\()lution within historic tinu-s. Most
ol the common species occur at mid-alti-
tudes in the mountains, where not much
carabid collecting was done until [']\'elyn
Gheesmans time, in the If).3()'s, and where
really extensiNC eolleetions of ('arabidae
have been made only recentK, by Dr.
Co-(\ssitt. the Se(llae(>ks. and other l^ishop
The Carabid Beetles of New Guinea • Darlington 145
Museum entomologists. Andrewes, during Beetles of New Guinea," Bull. Mus. Comp.
his work on Oriental Carabidae, did see a Zool. 126, No. 3, pp. 328-330).
few older specimens of Demctricki from In drawing descriptions in this genus I
New Guinea, including one or two of the have used "c." (circa, meaning approxi-
strikingly colored forms, but he refrained mately) even more often than usual, as a
from describing them; he did not know stratagem for saving space where I do not
what genus to put them in! So, I think think exact or detailed statements are use-
failure of earlier authors to describe New ful. I have also sometimes used it as a
Guinean species of Demctrida was a result warning that variation probably occurs al-
partly of the inaccessibility of the habitats though my material is too limited to show it.
of most common species, partly of com- A statement of my procedure in attacking
mendable caution on the part of taxonomists the particularly difficult problem presented
including H. E. Andrewes, and probably by the New Guinean Demetrida may be
partly just of chance. useful to future taxonomists. My method
Methods. My specific descriptions in has been to alternate between the general
Demetrida follow a special, slightly modi- and the particular, with first a general
fied form designed to characterize the sorting of individuals into apparent species
species adequately without wasting space, and preparation of a very preHminary key.
Characters covered in the generic Descrip- then drawing of detailed descriptions
tion are not repeated, but each specific species by species to determine characters
description begins with a statement that and variation, then preparation of an im-
the species shares the generic characters. In proved general classification and an im-
addition to the usual proportions, the ratio proved key, then further checking of details
of width of base of prothorax width of head and variation, and eventually (by a much
is given; it is especially useful in distin- longer process than this! ) preparation of a
guishing some species of Demetrida. The final key and descriptions emphasizing
headings Inner icings. Lower swface, and characters that have proved significant and
Leii,s are omitted; these subjects are suf- emphasizing variation, and last of all com-
ficiently covered in the Description of the pletion of introductory and explanatory ma-
genus. A special heading Claws is added terial, including the present statement. This
because number of claw teeth may prove is the general method that taxonomists use
to be diagnostic of some species. in classifying any unknown animals, but
Secondary sexual characters, especially the process has been much more complex
modifications of the S middle tibiae, have in Demetrida than usual. Specific problems
been examined carefully and used in char- have been numerous and difficult. In some
acterizing species. The tibiae are best seen cases I cannot be sure from available ma-
against an illuminated white background, terial whether differences in color, presence
To see a tibia clearly at the proper angle or absence of setae, or length of elytral
it is often necessary to straighten a middle spines are specific characters, cases of
leg, and this can usually be done without Mendelian dimorphism, or other individual
relaxing the specimen, by pulling the tibia variations. I have had to decide these
straight with a pin point and putting a cases arbitrarily, and my groupings of
minute drop of glue on the articulation to species are partly arbitrary. Tlie resulting
hold the straightened tibia in place. How- classification is at best an approximation,
ever, I have not examined the 6 copulatory Of course this is true of most classifications,
organs. This is a task for third-stage taxono- but I am more than usually conscious of
mists, far beyond what I have time to do the fact in this case,
now ( see Part I of my "The Carabid The question may be raised, why publish
146 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
a classification that is only a doubtful ap-
proximation? The answer is that it presents
an exciting situation in the only way that it
can be presented now. Further work on
Demetrido, including field work (which is
essential), requires some sort of classifica-
tion. In this and many other cases where
a classification is needed, w^e can only
follow what I think is a basic rule of the
trade: a taxonomist must do the best he
can with the available material in the avail-
able time. Actually, this imperfect treat-
ment of Dcmetrida may prove to be the
most important part of my taxonomic work
on New Guinean Carabidae.
The following Key to Species of Demetrida
of New Guinea is complicated and at some
points difficult to use. This is inevitable in
the case of an "exploding" group in which
some species are exceptionally variable and
others connected by intergrades. The key
is designed primarily for identification. It
is partly but not wholly phylogenetic:
species that are closely grouped in the key
are likely to be related but are not neces-
sarily so. A few dimorphic or exceptionally
variable species are run out at two different
points in the key, but I have had to limit
such multiple treatments. A few individuals
arc therefore unidentifiable by key char-
acters and have to be placed by comparison
of specimens. I might be able to construct
a multiple-tn-atmcnt kc-y that would iden-
tify every iiidixidual variant of eacli
Demetrida now known from New Guinea,
but the key would be impossibly complex
even lor present use and it would not take
care of new material, which will surely
include new variants of many species. The
key must be used with care and discretion.
Proportions must be calculated from inea-
sinements. Much variation must be allowed
for, more than I have been able to indicate
in detail. Alternatives must be tried when
specimens do not key out clearly. The test
of a key like this is whether it works rea-
sonably well in practice. First-time users
will probabK' Hud it \{'r\' difficult. Persons
who become more familiar with it will, I
hope, find shortcuts, in part suggested by
section headings inserted in brackets.
For comparisons of New Guinean De-
metrida with Australian species see espe-
cially Footnote .3 ( p. 143 ) and under D.
j)rima (p. 150).
Key to the Species of Demetrida of
Neav Guinea
[Pubescent]
1. Most of upper surface including sides of
head behind eyes plainly pubescent 2
- Surface not pubescent or (.scriata and
nuhicola only) pubescence very sparse,
fine, scarcely detectable 8
2. Pubescence short; elytra truncate or sinuate-
truncate at apex 3
- Pubescence long, sparse-pilose: elytra usu-
ally lobed or spined at apex ( scarcely so
in pallen.s) 6
3. Posterior-lateral prothoracic setae present,
at basal angles; smaller, length under 7.5
nmi (p. 149) aitape
- Posterior-lateral prothoracic setae absent;
size larger 4
4. Eyes more prominent, head nearly as wide
as prothorax ( width head/prothorax 0.98
and 0.96); 9 with 3 ( .^probably 2) apical
ventral setae each side; (length c. 9 mm)
( p. 149 ) goroka
- Eyes less prominent, head relatixely nar-
rower; apical ventral setae more numer-
ous 5
5. Color entirely brown; length usualK' more
than 9 nun (p. 150) priiiui
- Bicolored, head and prothorax brown,
elytra nearly black: length usually less
tliiui 9 mm (p. 151) __ fi/grZ/K'/ui/.v
(1 b^Ktral apices siiniate-truucate or we;ikl\'
lobed; color irreguLir p;ile brown (Y>.
151 ) pallcus
- Elytral apices spined 7
7. brown, el\ tra uitli pale speckles (p.
151 ) tcsscldtd
- Almost black, el\ tra faintly or not speckled
(p. b52 ) crcpciii
[I'UijtKi tiuncdic]
(S. l']|ytral apices oblicjuc^b' truncate or sinu;ite-
truucate (Figs. 97, 99) 9
- Elytral apices ;mgulate, toollied, spined,
or at le;ist subaugnhitely lobed (Fig. 102)
c. opposite ends of 2n(l striae or 2nd
iuter\als 15
9. TImHI and iisn;ill\ 5tli eblial intcrxals e;ieli
with se\('r;il st't;i-be;n ing punetuies; upper
suilaee with ;i little sp;irse, line, scarcely
(leteetiible jMibescence 10
The Carabid Beetles of New Guinea
Darlington
147
- Third intervals with 1 or 2 and 5th inter- 20.
vals without seta-bearing punctures; upper
surface without such pubescence; (color -
brown) 11 21.
10. Color entirely brown (p. 153) seriata
- Bicolored, reddish brown with base of
elytra black; (see also Description) (p.
153 ) mibicola
11. Large (over 10 mm) and prothorax
strongly narrowed in front and outer 22.
angles of elytra distinct, only slightly
blunted (p. 154) magna
- Either smaller or with prothorax differently
shaped or with outer elytral angles
rovmded 12
12. Prothorax wider than long at middle (by
measurement); posterior-lateral prothoracic 23.
setae usually present 13
- Prothorax longer than wide; posterior-
lateral prothoracic setae alisent 14 _
13. Larger (7-9.8 mm); outer angles of elytra
distinct although sometimes slightly blunted 24.
(p. 155) truncata
- Smaller (5.6-6.3 mm); outer angles of
elytra bhuited or rounded (p. 155) _^.- minor
14. Third elytral intervals 2-punctate; apical
ventral setae 3 each side in i , probably 25.
more in 9 (p. 156) std)temti.s
- Third elytral intervals 1-punctate; apical
ventral setae 1 each side in S , 2 in $ ( p.
156 ) tenuis
[Elytra ungulate, toothed, or spined]
15. Pronotum without posterior-lateral seta-
bearing punctures 16
- Pronotum with posterior-lateral seta-bear-
ing punctures at or near posterior angles --. 57
16. Color above brown (testaceous to piceous),
reddish, black, or bicolored black-and-
paler, but not in any part metallic 17
- Color above partly or wholly metallic
blue, blue-black, green, or purple, often
but not always bicolored 50
17. Not distinctly bicolored above, usually ±
uniform lirown, sometimes grading into og
darker brown or piceous on some parts _
of body ( some doubtful species are run
both ways in the key) 18 £7
- Sharply bicolored above, partly brown or _
reddish, partly (sometimes only broad
humeral areas) black 39 28.
18. Genae angulately or roundly prominent; _
(3rd intervals of elytra 3-punctate) 19 29.
- Genae oblique or weakly rounded, not
prominent 20
19. Elytral apices obtusely angulate (p.
157) tripuncta -
- Elytral apices acutely toothed ( p. 158 ) -^_.
_._ genicula
EKtral apices weakly lobed or angulate
with the angles blunted, obtuse, or right 21
EKtral apices acutely dentate or spined __ 23
Prothorax subcordate, wider than head and
much wider than long ( width/length c.
1.35); length 5.5-7.1 mm (p. 158) __ latangula
Prothorax subcjuadrate, usually (not always)
narrower than head and not or not much
wider than long; size larger 22
Prothorax slightly narrower (cf. descrip-
tions); i middle tibiae slightly bent-in
at apex; length 7.5-8.9 mm; (Papua) (p.
159 ) angulata
Prothorax slightly wider; $ middle tibiae
slightly bent-out at apex; length 8.5-9.2
mm; (West N. G. ) (p. 159) reversa
Median impressed line of pronotum fine;
(form as Fig. 103, very elongate; length c.
10-11 mm) (p. 160) kokoda
Median impressed line of pronotum
coarse 24
Prothorax at middle usually wider than or
equal to width of head, or only slightly
narrower 25
Prothorax much narrower ( usually by %o
or more) than width of head 33
Prothorax more cordate, usually wider (c.
% to 'h-i wider than long, with base often
wider than head) with sides more rounded
and more evenly rounded anteriorly, and
with wider margins, and 3rd elytral interval
with only two dorsal punctures, and color
entirely brown, and length usually less
than 7 mm 26
Either prothorax more quadrate and nar-
rower (less than Mi wider than long, with
base usually narrower than head) with
sides often but not always less evenly
rounded anteriorly and with narrower
margins, or 3rd elytral interval with more
than t\\Q dorsal punctures ( additional ones
sometimes smaller than the 2 primary
ones), or color darker (at least partly
blackish), or size larger 27
Elytral apices spined (p. 160) moda
Elytral apices acutely angulate or short-
toothed (p. 161) suhmoda
Prothorax with wider margins, ± cordate 28
Prothorax with narrower margins, quadrate
or more narrowly cordate 31
Smaller, length less than 8 mm 29
Larger, length 8 mm or more 30
Eyes more prominent; base of pronotum
less punctate; elytra usually with (some-
times faint) reticulate microsculpture (p.
162 ) hollandia
Eyes less prominent; base of pronotum
more punctate; elytra without reticulate
microsculpture (p. 162) toau
148 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
30. Third elytral interval 2-punctate (p. 163) _ 41.
similis
- Third elytral interxal with more than two -
punctures (p. 164) dupliccita
31. Smaller, length 6.7-7.6 mm; ± piceous; 42.
base of prothorax more punctate; 3rd
elytral interxals with more than two punc- -
tures (p. 164) suhpunctaia
- Larger, length 7.7-9.2 mm; browii; base 43.
of prothorax less punctate; 3rd elytral in-
tervals 2-punctate 32 _
32. Male middle tibiae tuberculate-serrate;
length 7.7-9.2 mm (p. 165) dohodura
- Male middle tibiae not tuberculate-serrate;
length c. 10.8 mm (p. 166) kiiin^a
[Protliorax much uanoucr than head]
33. Elytral 3rd intervals 2-punctate, and color
reddish brown \\ ith legs pale except knees
.sometimes dark, and length not over 10
mm 34
- Elytral 3rd intervals 3-punctate, or ( // 3rd
interval 2-punctate) color in part darker
with legs partly or wholly dark, or size
over 10 mm 36
34. Most of upper surface distinctly micro-
reticulate (p. 166) mafidii
- Microreticulation distinct (if at all) ouK
on elytra 35
35. Male middle tibiae tuberculate-serrate;
sides of prothorax usually more rounded-in
at apex (p. 167) forma
- Male middle tibiae bent-in at apex but not
tuberculate-serrate; sides of prothorax usu-
ally almost straight anteriorly (p. 167) recta
36. Very large, length 10.2-11.4 nun, and
basal angles of prothorax very prominent
(Fig. 105) (p. 168) L rex
- Smaller; l:)asa! angles of prothorax usualK
less promiui'iit -. 37
37. Legs red or brown, not darker llian disc
of elytra (p. 169) hitntnca
- Legs partly or wliolly dark _ 38
38. Third inter\als of elytra usually 3-pune-
tate; pronotum witlumt reticnkitc micro- _
sculpture (p. 169) .. . fumipcs ^
- Third intervals usually 2-punctate; pro-
notum ( h'glitly ) mierori'ticulate ( [). 170)
vclata
[Bieolorcd hut not mclalJic] 52.
39. Head and pronotum retldisli, cl\ tr;i en-
tirely dark or with only apices slightly -
reddish 10 r>:].
- Pattern not as described; elytra usually
(not always) bicolored 42 -
40. Lh Iral striae very lightly impressed; le.gs
l)lack (p. 171) ni^ri))e.s
- Elvtral striae well impressed; legs usually 54.
paler _ 41
44.
45.
46.
47.
48.
49.
[A/<
50.
El\tra usualK with reticulate microsculp-
tnre (p. 162) hoUandia
Elytra without reticulate microsculptiu'e
(see also couplet 29) (p. 162) wait
Prothorax wider (c. V-, or more wider than
long ) 43
Prothorax narrower ( c. long or longer than
wide ) 44
Elytra black with large connnon discal
area red (p. 171) dorsali.s
Elytra reddish brown with dark base
(western N. C;.; for individuals from
central and eastern N. C. see 'Notes imder
duplicata, p. 164) (p. 172) hasalis
Elytral apices angulatc: (color diverse,
see Description ) (p. 172 ) diversa
Ehtral apices spined (spines sometimes
short ) 45
Eyes abrupt (Fig. 107); (anterior angles
of protliorax more rounded than usual ) ( p.
173 ) vigil
Eyes normal, prominent but less abrupt 46
Head and pronotum black or piceous, or
at least darker than elytral disc 47
Head and pronotum red or brown 48
Elytra wholly brown, long-spined (p.
174) nigriceps
Elytra brown with darker humeri, shorter-
spined (p. 174) saidor
Elytra red or brown with base wholly
black, except suture sometimes red to base;
le.gs pale, sometimes with dark knees ( p.
175 ) ilivisa
EK tra with onl\- liiuntai dark; legs dark or
l)icolorcd 49
Dark humeral ari'as wider, usualK in-
chiding parts of 5th inter\als; lengtli 9..3-
10.8 mm (p. 175) humeralis
Dark humeral areas narrower; length 8.3-
9.4 mm (p. 169) _.. ftimi))cs
iaUic at least in jiart]
Broad ( bro;idcst i)i inctrida) : color en-
tircK bhic-black (p. 176) imitatrix
More slender; color not ;is described 51
ilc;id ;in(l prothor;ix red, eKtr;i entirely
green cxcipt sometimes jiurplisli posteri-
orK (p. 177) -- I iridipi-nni.\
(]()l(ii' I II it as dcsciibcd 52
Clolor entircK' green, .urecn ;uid black, or
gi'cen ;uKi purple 53
i^icolorcd, cKtrii in part icd or brown 54
Linger (9.2-10.8 mm); cncs more iibrupt;
eKtra lon.g-spined (p. 177) U'pida
Smaller (7.0-9.0 mm); eyes less ;ibrupt;
(Kli.i nsiialK sliorliT-spiiH d (p. 178) ...-
std)Iepida
llc;id :in(l j^roiiotum led or blown, not
I'l.iiiiK nu'lallic- 55
The Carabid Beetles of New Guinea • Darlingtun
149
- Pronotuni and sometimes also head metallic
green 56
55. Elytral apices angnlate (p. 172) __._ diversa
- Elytral apices short-spined (p. 179)
viridibasis
56. Elytral apices acutely annulate; most of
upper surface microreticulate (p. 166) iiiafulii
- Elytral apices long-spined; only elytra
(faintly) microreticulate (p. 179) sibil
{Posterior-lateral setae presciit]
57. Color hrown-piceous (head and pronotum
usually darker than elytra); prothoracic
margins with extra setae anteriorly (p.
180 ) seticollis
- Color wholly or mainly black, blue-black,
or green-black; prothoracic margins with-
out extra setae anteriorly 58
58. Legs red testaceous (p. 181) pallipes
- Legs dark 59
59. Elytra black with common discal area red
(p. 181) discoidalis
- Elytra metallic blue-black or green-black
without discal red area 60
60. Elytra short-spined ( p. 182 ) .— sedlacekorum
- Elytra long-spined (p. 182) hrandti
Demetr'ida aitape n. sp.
Description. With characters of genu.s;
form nearly as in following species {fS,oroka,
Fig. 97), slender, eyes prominent, elytral
apices obliquely subtruncate; entirely brown;
surface short-pubescent, without r(>ticulate
microsculpture but sparsely punctulate.
Flead 0.91 and 0.90 width prothorax; eyes
moderately prominent, genae shorter and
oblique. PwtJiora.x narrowly subcordate;
width/length 1.10 and 1.13; base/apex 1.36
and 1.34; base/head 0.94 and 0.91; sides
broadly rounded in anterior %, broadly
sinuate before right or slightly acute some-
times minutely blunted posterior angles;
margins rather wide in proportion to width
of insect, each with seta before middle
and at or just before basal angle; baso-
lateral areas slightly depressed and more
closely punctate than disc. Elytra long;
width elytra/prothorax 1.69 and 1.61; apices
slightly obliquely sinuate-truncate, with
outer angles moderately and sutural angles
narrowly rounded; striae deep, subpunctate
(or with sides of intervals slightly irregu-
lar); 3rd interval with c. 4 special seta-
bearing punctures (difficult to distinguish
amid other punctation). Claws with c. 6
teeth. Secondary sexiial characters: S tarsi
as genus; i middle tibiae tuberculate-ser-
rate ( c. 5 small tubercles ) ; 6 with 3 or 4,
$ 4 or 5 apical ventral setae each side.
Measurements (types only): length 5.8-
7.2; width 1.8-2.3 mm.
Types. Holotype $ (M.C.Z., Type No.
31,439) and 3 paratypes from Aitape, N-E.
N. G., Aug. 1944 (Darlington); 6 para-
types, Mt. Lamington, Papua (C. T.
McNamara, S. Australian Mus.).
Additional jnaterial. Papua: 1, Do-
bodura, Mar.-July 1944 (Darlington). N-E.
N. G.: 1, Erima, Astrolabe Bay, 1896 (Biro).
West N. G.: 1, Dojo [near Hollandia],
Apr. 1958 (G. den Hoed, Louwerens Coll.,
to Leiden Mus. eventually).
Measured specimens. The c^ holotype and
1 9 paratype.
Notes. The relatively small size and
presence of posterior-lateral prothoracic
setae distinguish this species from other
short-pubescent ones in New Guinea. The
individuals listed under Additional material
are slightly larger than the types and vary
slightly in form, but seem to be conspecific.
x\ll have setae at the posterior angles of
the prothorax as well as before middle,
and all are 9 9 with 5 setae each side last
ventral segment.
Demetrida goroka n. sp.
Description. With characters of genus;
form as in Figure 97, slender, eyes promi-
nent, elytral apices obliquely subtruncate;
reddish brown, elytra darker but not black,
appendages brown; surface short-pubes-
cent, without reticulate microsculpture but
sparsely irregularly punctate. Head 0.98
and 0.96 width prothorax; eyes rather
abruptly prominent, genae slightly shorter,
sinuate-oblique; front flattened, irregularly
impressed. Frothorax narrowly subcordate;
width length 1.04 and 1.02; base apex 1.26
and 1.38; base/head 0.86 and 0.91; sides
broadlv rounded in anterior %, broadly
150 BuUetin Museum of Comparative Zoology, Vol. 137, No. 1
sinuate before sharply acute posterior
angles; margins rather wide in proportion
to width of insect, each with seta slightly
before middle but none at base; baso-
lateral areas irregularly impressed, punc-
tate. Elytra long; width elytra /prothorax
1.69 and 1.68; apices slightly obliquely
sinuate-truncate, with outer angles blunted
or rounded, sutural angles narrowly
rounded; striae deep, irregularly subpunc-
tate; 3rd intervals each with 4 special
punctures in type, the punctures in part
obscured or absent in paratype. Chas with
6 or 7 teeth. Secondary sexual characters:
$ unknown; 9 with 3 apical ventral setae
each side in both specimens. Measure-
ments: length ± 9.0; width 2.8 mm.
Types. Holotype 9 (Bishop Mus.) and
1 9 "paratype (M.C.Z., Type No. 31,440)
both from Goroka, N-E. N. G., 1500 m.
May 22, 1961 (J. L. & M. Gressitt), taken
in light trap.
Notes. D. <:oro1<a superficially resembles
prima and nin,ripennis (below) but has
wider head \\\\\\ more prominent eyes,
slightly narrower prothorax ( with more dis-
tinctly lobed base), and only 3 apical ventral
setae each side in 9 (probably only 2 in $),
while 9 9 of prima and nigripennis have
5 or 6 such setae each side.
Demetrida prima n. sp.
Descri])tion. With characters of genus;
form nearly as in preceding species (goroka.
Fig. 97), slender, but with eyes less promi-
nent than in goroka, elytral apices obliquely
subtruncate; reddish brown; surface short-
pubescent, without reticulate microsculp-
ture but sparsely punctulate. Head 0.91
and 0.87 width prothorax; eyes less promi-
nent than usual in genus, genae long-
{)bli((uc. Prothorax narrowly subcordate;
width/length 1.03 and 1.08; base/apex 1.24
and 1.11; base/head 0.96 and 0.93; sides
weakly arcuate anteriorly, broadly sinuate
before right or slightK' acute well defined
basal angles; margins moderate, (\ich ap-
parently with special seta-bearing punc-
ture near middle of length but not at base
(these setae and punctures difficult to
distinguish amid general pubescence); baso-
lateral impressions punctate. Elytra long;
width elytra prothorax 1.64 and 1.53; apices
truncate or weakly sinuate-truncate, with
outer angles moderately and sutural angles
more narrowly rounded; striae deep, ir-
regularly punctate; 3rd interval with ap-
parently 2-5 special seta-bearing punctures
( sometimes difficult to distinguish amid
other punctation ). Clans with 5 or 6 teeth.
Secondary sexual characters: 6 unknown;
9 with c. 5 apical ventral setae each side.
Measurements: length 9.0-9.8; width 2.8-
3.0 mm.
Types. Holotvpe 9 (Bishop Mus.) from
Wau, Morobe Dist., N-E. N. G., 1200 m.
Mar. 23, 1963 (Sedlacek), in mercury vapor
light trap. Additional (99) paratypes
from N-E. N. G. as follows: 1, Maprick,
160 m, Dec. 29, 1959-Jan. 17, 1960 (T. C.
Maa, now in M.C.Z., Type No. 31,441); 1,
Torricelli Mts., Siaute, sea level, Nov. 9-17,
1958 (W. W. Brandt, Bishop Mus.); 1,
Mumeng, 600 m. Mar. 9, 1962 (Sedlacek).
Measured specimens. The 9 holotype and
9 paratype from Mumeng.
Notes. Among New Guinean Demetrida,
prima should be easily known by form,
including form of elytral apices, rather large
size, nearly uniform reddish brown color,
and short-pubescent surface.
Superficially prima resembles some Aus-
tralian species of Demetrida. For example
it is somewhat similar in form to grandis
(Chaudoir) of southern Australia but has
shorter antcnmae, smaller eyes, less promi-
nent genae, prothorax narrower anteriorly
with narrower margins, and elytra uni-
formly brown (not strip(>d as in grandis).
D. prima also somewhat resembles con-
stricticeps (Sloane) of southwestern Aus-
tralia in lonn and is similar in color, but
]nima has shorter antennae, much less
prominent genae, less strongU' simiate sides
ol prothorax, and dilleis in other details.
And prima differs Ironi l)()th the Aus-
The Carabid Beetles of New Guinea • Darlinaton
151
tralian species named and from all other
Australian species known to me in amount
and character of pubescence.
Demetrida nigripenn'is n. sp.
Description. See Plate 1, figure I; with
characters of genus; form, elytral apices,
pubescence, punctation, and other asexual
characters c. same as in preceding species
(prima), but color brownish red with
elytra black or nearly so, and size smaller.
Head 0.88 and 0.89 width prothorax. Pro-
thorax: width/length 1.06 and 1.07; base/
apex 1.20 and 1.18; base/head 0.90 and
0.98. Elytra: width elytra/prothorax 1.56
and 1.64; 3rd intervals with apparently 1-4
principal seta-bearing punctures (difficult
to distinguish). Secondary sexual char-
acters: S tarsi as genus; 6 middle tibiae
weakly 2-emarginate on inner edge near
apex; S with c. 4, 9 c. 6 apical ventral
setae each side. Measurements: length
8.3-9.0; width 2.5-2.8 mm.
Types. Holotype S (Louwerens Coll.,
eventually to Leiden Mus.) and 1 S para-
type (M.C.Z., Type No. 31,442) from Dojo
[near Hollandia], West N. G., Apr. 1958
(G. den Hoed); and 1 9 paratype, Hol-
landia, May 1945 (B. Malkin, U.S.N.M.).
Measured specimens. The i holotype and
9 paratype.
Notes. This may prove to be a geographic
form of the preceding species ( prima ) , but
more material of both sexes from more
localities is needed to clarify the relation-
ship. The form of the c^ middle tibiae is
unique among known members of the
genus.
Demetrida pailens n. sp.
Description. See Plate 1, figure II; with
characters of genus; eyes prominent, pro-
thorax small, cordate-quadrate, and elytral
apices sinuate-subtruncate and usually sub-
angulate c. opposite ends 2nd striae; color
irregular testaceous brown, elytra irregu-
larly tesselated with small paler spots; sur-
face long-pubescent, without reticulate mi-
crosculpture, punctate as described below.
Head 1.10 and 1.08 width prothorax; eyes
prominent, genae short-oblique. Prothorax:
width/length 1.09 and 1.03; base/apex 1.36
and 1.38; base/head 0.85 and 0.91; side
margins moderate, entirely fringed with
long setae; disc irregularly sparsely punc-
tate. Elytra: width elytra/prothorax 2.07
and 2.17; apices usually as figured, sub-
angulate or weakly lobed (simply sinuate-
truncate in 1 specimen), outer angles
broadly and sutural angles less broadly
rounded; striae moderately impressed, in
part slightly interrupted, irregularly sub-
punctate; intervals all with series of coarse
seta-bearing punctures among which special
dorsal punctures are not distinguishable.
Claws with 7 or 8 teeth. Secondary sexual
characters: 6 tarsi as genus; S middle
tibiae tuberculate-serrate (c. 6 small tuber-
cles); c^ with c. 4 apical ventral setae each
side; 9 unknown. Measurements: length
8-9; width 2.9-3.3 mm.
Types. Holotype c5 (Leiden Mus.) and
4 $ S paratypes (2 in M.C.Z., Type No.
31,443) all from Moss Forest Camp (Snow
Mts.), West N. G., 2800 m, Oct. 9-Nov. 5,
1938 (Toxopeus).
Measured specimens. The $ holotype and
1 9 paratype.
Notes. This very distinct species occurs
at a higher altitude than any other
Demetrida known to me. The coloration,
which superficially resembles that of some
high-altitude Agonini (some Mactdagonum),
suggests that the insect lives in grass, al-
though the specimens were taken at "Moss
Forest Camp."
The (slight) variation in form of elytral
apices is one of many examples of individ-
ual variation in this remarkably variable
genus.
Demetrida fesselata n. sp.
Description. With characters of genus;
form ( Fig. 98 ) c. average, with eyes promi-
nent, prothorax small, elytra spined; color
irregular dark reddish brown, elytra with
152 BiiUetin Museum of Comparative Zoology, Vol. 137, No. 1
numerous small pale flecks forming rows
most conspicuous on ( but not confined to )
odd intervals, legs pale; surface sparsely
long-pubescent, without reticulate micro-
sculptme, irregular but scarcely punctate
except for punctures (variable in size) from
which hairs rise. Head 1.16 and 1.08 width
prothorax; eyes prominent, genae nearly as
long as eyes, oblique. Prothorax small, nar-
ro\N']y cordate-subquadrate; width length
1.03 and 1.07; base apex 1.31 and 1.35;
base head 0.81 and 0.88; sides weakly
rounded anteriorly, often subangulate near
middle of length; side margins moderate,
irregularly fringed for entire length with
long setae; baso-lateral depressions poorly
defined, irregular but scarcely punctate.
Elytra: width elytra prothorax 2.28 and
2.23; apices spined, outer angles rounded
or obtusely blunted (variable), sutural
angles obtusely blunted; striae lightly im-
pressed, in part interrupted or reduced to
rows of punctures; intervals flat but ir-
regular, odd intervals with series of seta-
bearing punctures of moderate size, each
puncture usually on posterior side of a
broad low tubercle. Claws with c. 6-8 teeth.
Secondary sexual characters: 6 tarsi as
genus; i middle tibiae tuberculate-serrate
(c. 5 widely spaced small tubercles); i
with 3 (or more?), 9 5 or 6 apical ventral
setae each side. Measurements: length 8.7-
9.8; width 3.0-3.4 mm.
Types. Holotype $ (Bishop Mus.) and
1 9 paratype (M.C.Z., Type No. 31,444)
from Mt. Kaindi, N-E. N. G., 2350 in, Jan.
10 and June 9, 1962 (Sedlaceks), "the
paratype taken in mercury xapor light
trap; and 1 additional paratype i without
head, same locality, 2400 m, Jan. 28, 1963
(Sedlacek); 3 paratypes, Wau, 2400 m, Jan.
9-12, 1962 (Sedlaceks); 2 paratypes, 32 km
S. of Wau, Bulldog Rd., 2850 m. May 29-
30, 1962 (Sedlacek), light trap.
Additional material. Papua: 1, Mt. Tafa,
8500 ft. (c. 2600 m), Mar. 1934 (Cheesman).
N-E. N. (;.: 1 ,5, Edie Creek, 14 km S\A'
of \Yau, 2000 m, Fvh. 13, 1962 (S(>dlacek);
1 9 , Enarotadi, 2000 m, Aug. 1962 ( Sed-
lacek). West N. G.: I $, Swart Valley,
\V. ridge 1800-2000 m, Nov. 19, 1958
( Gressitt ) .
Measured specimens. The i from Edie
Creek and the 9 holotype, figures given
in this order.
Notes. The specimens before me vary not
only in size and color but also in fonn of
outer-apical elytral angles, depth of striae,
presence or absence of low rounded tuber-
cles on odd elytral intervals, and in other
ways. Some of this variation is surely in-
dividual, but some may be geographic.
Only additional series from several localities
can decide this.
Demefrida crepera n. sp.
Description. Form and characters c. as in
preceding species ( fesselata ) except color
piceous or slightly reddish piceous without
distinct pale flecks on elytra. Head 1.04
and 1.05 width prothorax. Prothorax:
width length 1.08 and 1.06; base apex 1.30
and 1.25; base head 0.85 and 0.85. Elytra:
width elytra prothorax 2.05 and 2.12; sculp-
ture somewhat variable but in general like
that in preceding species (tesselata), in
\\'hich the sculpture varies too. Secondary
sexual characters: 6 tarsi as genus; 6 mid-
dle tibiae tuberculate-serrate ( c. 6 tuber-
cles); 6 with c. 4, 9 c. 6 apical \entral
setae each side. Measurements: length 9.5-
10.4; width 3.1-3.5 mm.
Types. Holotype $ (A.M.N.H.) and 6
paratypes (2 in M.C.Z., Type No. 31,445)
from N. slope, Mt. l^ayman, Maneau Rge.,
Papua (the h()lot\pe and 4 parat\pes at
"No. 4,"' 2230 m, Ma>- 19-June 19, 1953; 1
paratype, same data except June 1-7; 1
paratype same except "No. 5," 1550 m. lune
30-Ju'lv 13) (all specimens collected 1953
by Ceoffrey M. Tate).
Additional material. N-E. N. G.: 1, "No.
10," Purosa Camp, Okapa area, 1950 m,
Sept. 29, 1959; 1, "No. 6," Pengagl Camp,
east slopes Mt. W'ilhelm, 2770 m, July 3,
1959 (both specimens Sixth Archbold Exp.,
L. 1. Hrass. A. M.N. 11.).
The Carabid Beetles of New Guinea • Darlington 153
Measured specimens. The S holotype and cles widely spaced ) ; S with 3, 9 4-6
1 5 paratype. apical ventral setae each side. Measure-
Notes. This form is apparently a geo- mcnts: length 5.6-7.6; width 2.0-2.8 mm.
graphic representative (perhaps eventually Types. Holotype i (Bishop Mus.) and
to be considered a subspecies) of the pre- 4 paratypes (2 in M.C.Z., Type No. 31,446)
ceding species (tesseJata) but is almost from Eramboe, 80 km ex Merauke, West
black rather than brown, is not distinctly N. G., holotype Feb. 1, paratypes Jan. 29,
pale-speckled, is slightly larger, and differs Feb. 5, 1960 (T. C. Maa).
slightly in proportions, especially in having Additional material Papua: 1 £ , Aroa
relatively narrower elytra. Estate, W. of Redscar Bay, 1 m, Sept. 29,
All individuals of the type series have 1958 (Gressitt); 1 $ , Bisianumu, E. of Port
moth scales stuck to them, indicating that Moresby, 500 m, Sept. 23, 1955 (Gressitt);
they were taken in light traps. 1, Daradae nr. Javarere, Musgrove R., 100
m?, Oct. 2, 1958 (Gressitt); 4, Mt. Laming-
Demetrida seriata n. sp. ton, 1300-1500 ft. {c. 400-460 m) (C. T.
Description. With characters of genus; McNamara, S. Australian Mus.).
form c. average, with prominent eyes, sub- Measured specimens. The 6 holotype and
cordate prothorax, elytra with sinuate- 1 ? paratype.
truncate apices and usually slightly nar- ^otes. This species is characterized by
rowed to\N'ard base; color brown or relatively numerous dorsal elytral seta-
testaceous; surface not obviously pubescent bearing punctures. Notable also is pres-
( actually very sparsely and inconspicuously ence of a little sparse, inconspicuous pubes-
so); reticulate microsculpture present (faint) cence (not visible on sides of head behind
only on elytra. Head 1.06 and 0.98 (some- eyes) and of several weak outward-directed
times less) width prothorax; eyes promi- hairs on margins of prothorax near anterior
nent, genae short-oblique, not prominent, angles. The species is very distinct and
Prothorax narrowly subcordate; \\'idth/ probably ranges over the whole length of
length 1.06 and 1.04 (wider in some speci- New Guinea although it has been found
mens); base/apex 1.32 and 1.30; base/head thus far only in two widely separated areas
0.87 and 0.87; sides broadly rather weakly near opposite ends of the island,
arcuate in anterior %, broadly sinuate before
c. right posterior angles; each side with Demetnda nub/co/a n. sp.
seta before middle and at posterior angle Description. See Plate 1, figure III; with
(all specimens) and additional weaker setae characters of genus; head, prothorax, and
directed more to side than upward near posterior part of elytra dark red, basal %
anterior angle; most of disc virtually im- of elytra black with black color extending
punctate. Elytra: width elytra /prothorax farther back at sides than at suture ( suture
1.92 and 1.97; apices obliquely sinuate- narrowly red), lower surface red (yellow-
truncate, with both outer and sutural angles ish on abdomen ) with metastema and con-
rounded or blunted; striae impressed, tiguous parts of epipleurae dark, femora and
scarcely punctulate; intervals convex, ir- outer edges of tibiae dark, tarsi and an-
regularly sparsely punctulate, 3rd with usu- tennae reddish yellow; not obviously pubes-
ally 6 ( sometimes fewer ) larger seta-bear- cent but with some sparse very inconspicu-
ing punctures, and 3 or 4 similar punctures ous hairs; reticulate microsculpture absent,
usually present on 5th intervals. Claws Head 1.06 width prothorax; eyes prominent,
with c. 5 teeth. Secondary sexual char- genae shorter, obliquely rounded into neck;
acters: S tarsi as genus; 6 middle tibiae front slightly convex, \\'ith 2 impressions
weakly tuberculate-serrate (c. 6 small tuber- anteriorly ( as usual in genus ) and ir-
154 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
regularly slightly impressed at middle. Pro-
thorax siibquadrate with base slightly
broader and apex narrower than usual;
width/length 1.03; base/apex 1.48; base/
head 0.92; sides weakly arcuate for much of
length, subangulate at median-lateral setae,
strongly sinuate before slightly acute promi-
nent posterior angles; margins moderate,
each with seta-bearing puncture c. % from
apex and at basal angle, and with several
much finer hairs directed laterally near an-
terior angles; disc moderately (not strongly)
convex, baso-lateral impressions present
but irregular, subpunctate. Elytra: width
elytra/prothorax 2.11; apices sinuate-trun-
cate, outer angles broadly rounded, sutural
angles blunted; striae well impressed, finely
punctulate; intervals convex, 3rd with c.
7 and 5th with 4 or 5 seta-bearing punc-
tures. Claws with c. 5 teeth. Secondary
sexual characters: i unknown; 9 with 4
apical ventral setae each side. Measure-
ments: length 7.8; width 2.8 mm.
Type. Holotype $ (Leiden Mus.) from
Lower Mist Camp [Snow Mts.], West
N. G., 1550 m, Jan. 31, 1939 (Toxopeus);
the type is unique.
Notes. This distinct species is the only
known Demetrida that combines unarmed
elytral apices with dual (black and red)
coloration. The sparse, very inconspicuous
pubescence and the extra seta-bearing
punctures of 3rd and 5th elytral intervals
are noteworthy too. The form of elytral
apices and the character of pubescence and
setae suggest a relationship with seriata,
but nuhicola is specifically distinct not only
in color but also in form of prothorax.
Demetrida magna n. sp.
Description. With characters of genus;
form large, slender, with large eyes, long-
quadrate or trapezoidal [)r()tliorax, elytra
sinuate-truncate at apex; color entirely red-
dish brown; surface not jMibescent, reticu-
late microsculpturc distinct (and sliglitly
transverse) only on elytra, punctation as
described below. Head 0.94 and 0.98
width prothorax; eyes prominent, genae
short and oblique. Prothorax long, sub-
parallel or trapezoidal; width length 0.98
and 1.00; base/apex 1.46 and 1.58; base/
head 1.01 and 1.01; sides broadly arcuate
anteriorly, usually broadly sinuate before
right or slightly acute usually blunted pos-
terior angles; margins moderate, each with
seta at basal angle and before middle; disc
fainth' or not punctulate except irregularly
subpunctate in baso-lateral depressions.
Elytra long; width elytra prothorax 1.92
and — ( elytra of 2nd specimen too spread
for measurement ) ; apices obliquely sinuate-
truncate, outer angles well defined (some-
times slightly blunted), sutural angles nar-
rowly rounded or blunted; striae impressed,
faintly punctulate; intervals convex, sparsely
punctulate, 3rd with 2 seta-bearing punc-
tures (present in all specimens but varying
in position). Cdows with c. 7 or 8 teeth.
Secondary sexual characters: S tarsi as
genus; $ middle tibiae weakly tuberculate-
serrate; £ with 3, 4, or 5 (number some-
times unsymmetric), 9 c. 6 apical ventral
setae each side. Measurements: length
10.3-12.0; width 3.3-4.2 mm.
Types. Holotype S (Bishop Mus.) from
Finschhafen, Huon Pen., N-E. N. G., 20-
150 m, Apr. 15, 1963 (Sedlacek). Para-
types as follows: N-E. N. G.: 2 ( ,$ 9 ),
Pindiu, Huon Pen., 870-1300 m, Apr. 20,
21-22, 1963 (Sedlacek, M.C.Z., Type No.
31,447); 1 9 , Adalbert Mts., Wanuma, 800-
1000 m, Oct. 25, 1958 (Gressitt); 1, Mark-
ham R., 10 m, Jan. 18, 1961 (Sedlaceks).
Papua: 1, Kokoda, 1200 ft. (366 m). May
1933 (Cheesman); 1, Owen Stanlev Rge.,
Goilala, Loloipa, Feb. 1-15, 1958 (w. W.
Brandt. Bishop Mus.); 1, Mt. Lamington,
1300-1500 ft. {c. 400-460 m) (C. T.
McNamara, S. Australian Mus.). West
N. G. : 1 ^ , Guega, \\\ of Swart Valley,
1200 m, Nov. 15, i958 (Gressitt).
Measured s])ecimens. The i holotype and
9 paratype from Adalbert Mts.
Notes. Comparative characters of mapia
are gi\(>n in tlie preceding Key. The species
The Carabid Beetles of New Guinea • Darlington 155
appears to be widely distributed at low
altitudes in New Guinea, but not common.
Demetrida truncata n. sp.
Description. With characters of genus;
form c. average, but variable; entirely
reddish brown; not pubescent, reticulate
microsculpture present ( but light and vari-
able) only on elytra, and surface not or
not much punctulate. Head 0.96 and 0.91
width prothorax; eyes moderately promi-
nent, genae much shorter, oblique. Pro-
thorax rather long, variable in shape (nar-
rowly subcordate to trapezoidal); width/
length 1.07 and 1.05; base apex 1.37 and
1.38; base/head 0.95 and 1.02; sides vari-
ably arcuate anteriorly, broadly sinuate be-
fore right or slightly acute blunted or well
defined posterior angles; margins moderate,
each usually with seta at or near basal
angle and before middle (but see Notes
below); disc smooth at middle, slightly
wrinkled or subpunctate at base and sides.
Elytra long; width elytra prothorax 1.80
and 1.78; apices obliquely sinuate-truncate,
outer angles well defined (c. right but
shghtly variable), sutural angles narrowly
rounded; striae impressed, punctulate; in-
tervals ± punctulate, often with an irregu-
lar row of small punctures near middle each
interval, 3rd with 2 dorsal punctures in all
specimens. Claws with 7 or 8 teeth. Sec-
ondary sexual characters: $ tarsi as genus;
S middle tibiae finely tuberculate-serrate
(about 9 slight tubercles); S with 3 or 4,
9 5 or 6 apical ventral setae each side.
Measurements: length 7.0-9.8; width 2.5-
3.5 mm.
Types. Holotype $ (Bishop Mus.) from
Wau, Morobe Dist., N-E. N. G., 1150 m,
Oct. 16, 1961 (Sedlacek); and paratypes as
follows. N-E. N. G.: I 6, Wau, Mt. Mis-
sim, 880-1050 m, Feb. 8-9, 1963 (Sedlacek);
1 9 , Busu R., E. of Lae, 100 m, Sept. 14,
1955 (Gressitt); 1 9, Finschhafen, Huon
Pen., 180 m, Apr. 16, 1963 (Sedlacek); 1 i ,
Torricelli Mts., Mobitei, 750 m, Feb. 28-
Mar. 4, 1959 (W. W. Brandt, Bishop Mus.).
West N. G.: 1 9 , Hollandia, Nov. 21, 1944
(H. Hoogstraal, M.C.Z.); 1 9 , Wans, S. of
Hollandia, 450-500 m, Aug. 16-23, 1959
(T. C. Maa, Bishop Mus.); 1 9, Jutefa
Bay, Pim, sea level-100 ft. (30 m), Feb.
1936 ( Cheesman ) ; 1 $ , mountain slope
above Bernhard Camp, 750 m, Mar. 1939
(Toxopeus). (Some paratypes in M.C.Z.,
Type No. 31,448.)
Additional material N-E. N. G.: 1 9,
Finisterre Rge., Saidor: Aiyawa Village,
June 16-23, 1958 (W. W. Brandt, Bishop
Mus.). West N. G.: 1 teneral 9, Hol-
landia, 250 ft. (c. 75 m), Nov. 3, 1944
(H. Hoogstraal, M.C.Z.); 1 9, Camp 1,
Mt. Xok, Waigeu Is., 2500 ft. (c. 760 m).
May 1938 (Cheesman).
Measured specimens. The i holotype and
the 9 paratype from Busu R.
Notes. This species (if it is all one
species) is widely distributed at low alti-
tudes in central and western New Guinea.
It is not recorded in Papua and may be
replaced there by the following species
(minor). Much more material from many
localities is needed to establish the specific
limits and geographic variation of these
forms.
Although truncata usually has a seta-
bearing puncture at or near each posterior
prothoracic angle, the individual from
Jutefa Bay has a well developed seta on
the right but no trace of seta or puncture
on the left, and seta and puncture are
lacking on both sides in the individual from
the Finisterre Range. The types and other
specimens listed above vary in other ways
the significance of which cannot be deter-
mined without more material. For example,
the prothorax is narrowly subcordate or
subquadrate in most of the types while
the individual from Waigeu Is. has the
prothorax strikingly trapezoidal, but the
extremes are connected by intermediates.
Demetrida minor n. sp.
Description. With characters of genus;
form as in truncata except outer apical
elytral angles c. rounded; reddish brown;
not pubescent, reticulate microsculpture in-
156 BuUetin Musciiiu of ConijMirafivc Zoology. Vol. 137. No. 1
distinct or lightly indicated on elytra, sur-
face very little punctulate. Head 0.89 and
1.02 width prothorax; eyes prominent,
genae short and oblique. Frothorax quad-
rate-subcordate; width/length 1.17 and 1.08;
base/apex 1.52 and 1.44; base/ head 1.03
and 0.97; sides broadly rounded in anterior
■")4, broadly sinuate before right-obtuse,
slightly blunted basal angles; margins mod-
erate, each with seta at base and before
middle (posterior seta-bearing puncture
present on both sides in all specimens);
disc scarcely punctate. Elytra: width
elytra/prothorax 1.71 and 2.02; apices
obliquely sinuate-truncate, outer angles
blunted or rounded, sutural angles narrowly
rounded; striae impressed, faintly or not
punctulate; intervals sparsely inconspicu-
ously punctulate, 3rd with 2 dorsal punc-
tures. Claics with c. 5 teeth. Secondary
sexual characters: i tarsi as genus; c^
middle tibiae weakly tuberculate-serrate;
S with 2, 9 4 apical ventral setae each
side. Measurements: length 5.6-6.3; width
2.0-2.2 mm.
Types. Holotype 6 (M.C.Z., Type No.
31,449) and 1 S paratype from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
1 9 paratype from Brown R., Papua, 5 m,
Oct. 23, 1960 (Gressitt).
Measured specimens. The A holotype and
9 paratype.
Notes. This species is probably related to
truncata but is smaller, with more rounded
outer apical elytral angles and fewer apical
vc^ntral setae.
Demetrida subfenuis n. sp.
Descriplion. With characters of genus;
form (Fig. 99) c. as in truncata and minor
but more slender; reddish brown; not
pubescent, reticulate microsculpture at most
faintly indicated on elytra, surface not
Diueli puiictiilate. Head 1.09 and 1.04
width prothorax; eyes proiniiiciit. genae
short and oblicjue, not jiioniinent. Pro-
tliorax el()ngate-su1)(ina(hal('; width length
0.89 and 0.94; base apex 1 .,36 and 1.40;
base/head 0.88 and 0.96; sides weakly
arcuate in much of length, broadly sinuate
well before c. right, scarcely blunted basal
angles; margins narrow, each with seta-
bearing puncture before middle but none
at base. Elytra: width elytra prothorax
2.08 and — ( elytra of 2nd specimen too
spread to measure); apices obliquely sinu-
ate-truncate, with outer angles obtuse and
slightly blunted or narrowly rounded and
sutural angles blunted; striae impressed,
faintly punctulate; intervals slightly punc-
tulate, 3rd with 2 dorsal punctures. Claics
with 5 or 6 teeth. Secondary sexual char-
acters: 6 tarsi as genus; i middle tibiae ±
bent-in and weakly tuberculate-serrate; S
with 3 apical ventral setae each side (both
sides both specimens); 9 unknown. Mea-
surements: length c. 7.0; w idth c. 2.3 mm.
Types. Holotype c5 (Bishop Mus.) from
Wum, Upper Jimmi Valley, N-E. N. G.,
840 m, July 17, 1955 (Gressitt); and 1
broken i paratype (M.G.Z., Tvpe No.
31,450), vie. Hollandia, West N. G., July-
Sept. 1944 (Darlington).
Notes. D. suhlcnuis resembles minor
(above) but is much narrower and lacks
posterior-latt>raI prothoracic setae, which
are present in minor.
Demetrida tenuis n. sp.
Dcscri})tion. With characters of genus;
form (Fig. 100) c. as in preceding species
(suhtcntii.s) but even more slender; brownish
led, legs slightly paler; not pubescent,
reticulate microsculpture visible (but very
light) only on elytra, surface not much
punctulate. IIc(ul 1.09 and 1.07 width pro-
thorax; eyes large, moderatcK' prominent,
genae much short(M-, oblicpie but conxexly
prominent; Iront carinate at middle anteri-
orl\- (all sj)i'eimens). ProtJiorax elongate-
(juadrate; width length 0.84 and 0.93; base/
apex 1.21 and 1.19; base head 0.87 and 0.83;
sides very wcakK' irregularK' anuulate, verv
broadly sinuate belore right or slightly
obtuse but well deliiicd basal angles; mar-
gins rati It r narrow, each with seta-bear-
The Carabid Beetles of New Guinea
Darlington
157
ing puncture before middle but without
posterior seta or puncture; disc faintly
pinictulate, wrinkled or subpunctate in
baso-lateral areas. Elytra: width elytra,
prothorax 2.36 and 2.32; apices sinu-
ate-truncate, outer and sutural angles ±
rounded; striae impressed, punctulate; in-
tervals slightly convex, faintly sparsely
punctulate, 3rd with 1 dorsal puncture, c.
Vi or less from apex (both sides all ex-
amples). Claws with c. 4 teeth. Secondary
sexual characters: 6 tarsi as genus; 6
middle tibiae tuberculate-serrate (4 tuber-
cles); S with 1, 9 2 apical ventral setae
each side. Measurements: length c. 6.0-
6.5; width 2.0-2.3 mm.
Types. Holotype $ ( Bishop Mus. ) from
Aroa Estate, W. of Redscar Bay, Papua,
1 m, Sept. 29, 1958 (Gressitt);' and 1 9
paratype (M.C.Z., Type No. 31,451) from
Owen Stanley Rge., Papua, Goilala: Lo-
loipa, Feb. 1-15, 1958 (W. W. Brandt); 1
c5 paratvpe, Brown R., 20 km N. of Port
Moresby, Apr. 29, 1960 (C. W. O'Brien,
Bishop Mus.).
Measured specimens. The 9 holotype and
9 paratype.
Notes. D. tenuis is characterized by small
size, very narrow form especially of pro-
thorax, carination of front, 1-punctate 3rd
elytral intervals, and small number of setae
of apical ventral segment. The last three
characters are unique among New Guinean
Demetrida, but some Australian species
have 1-punctate 3rd intervals, as indicated
in Notes under the genus and in Footnote 3
(p. 143).
Demetrida tripuncfa n. sp.
Description. With characters of genus;
form c. average, except genae angulate,
elytral apices obtusely angulate; reddish
brown; not pubescent, reticulate micro-
sculpture distinct ( but light ) only on elytra,
surface not much punctulate. Head 1.06
and 0.94 width prothorax; eyes large, promi-
nent (slightly variable), genae shghtly
shorter than eyes, subangulately prominent;
front flattened and irregularly impressed
anteriorly. Prothorax subquadrate; width/
length 1.04 and 1.06; base/apex 1.28 and
1.14; base head 0.80 and 0.87; sides
broadly irregularly arcuate in c. anterior %,
broadly sinuate before c. right but blunted
posterior angles; margins moderate, each
with seta before middle but none at
posterior angle; surface punctate-wrinkled
in baso-lateral impressions and margins.
Elytra: width elytra/prothorax — (elytra
spread) and 1.91; apices obtusely angulate,
with outer angles right or slightly obtuse,
sutural angles obtuse; striae impressed,
sometimes finely punctulate; intervals con-
vex, sparsely punctulate, 3rd with 3 dorsal
punctures (all specimens). Claws with c.
6 long teeth and sometimes an additional
minute one. Secondary sexual characters:
5 tarsi as genus; 6 middle tibiae slightly
bent-in near apex but not tuberculate-ser-
rate; i with apparently 3, 9 6-8 apical
ventral setae each side. Measurements:
length c. 8-9; width 3.0-3.3 mm.
Type. Holotype 9 (M.C.Z., Type No.
31,452) from Hollandia, West N. G., Nov.
21, 1944 (Hoogstraal).
Additional material. Papua: 1 S , Ori-
omo Govt. Sta., W. District, Oct. 26-28,
1960 (Gressitt), taken in Malaise trap; 1
9 , Brown R., 5 m, Oct. 23, 1960 (Gressitt),
taken on palm. N-E. N. G.: 1 9, Bulolo,
730 m, Aug. 15, 1956 (E. J. Ford, Jr.,
Bishop Mus.), taken in light trap.
Measured specimens. The 6 from Papua
and the 9 holotype, in this order.
Notes. The subangulate genae distin-
guish this species from all the preceding
ones except tenuis, which is very different
in many ways (see preceding Key to Species
of Demetrida).
The four specimens listed above agree in
a general way and in such important char-
acters as prominence of genae, 3-punctate
3rd intervals, and obtusely angulate elytral
apices, but they are from scattered localities
and they differ in many details. The single
6 is teneral and warped so that width of
158 Bulletin Miiscinii of Comparative Zoology, Vol. 137, No. 1
elytra cannot be measured, and some other
characters are difficult to see. More ma-
terial is needed to show whether all these
specimens really are conspecific.
Demefrida genicula n. sp.
Description. With characters of genus;
form (Fig. 101) as in preceding species
( tripuncta ) but elytral apices acutely
toothed; reddish brown; not pubescent,
reticulate microsculpture distinct (but light)
only on elytra, surface not much punctulate.
Head 1.08 and 1.03 width prothorax; eyes
prominent, genae subangulately prominent;
front flattened and irregularly slightly im-
pressed anteriorly. Prothorax quadrate-
subcordate; width length 0.97 and 1.04;
base/apex 1.24 and 1.19; base/head 0.(S3
and 0.S3; sides irregularly weakly arcuate in
anterior 'n, strongly sinuate before right or
slightly acute slightly blunted posterior
angles; margins moderate, each with seta-
bearing puncture just before middle but
none at base; disc weakly strigulose or sub-
punctate especially laterally. Elytra: width
elytra/prothorax 1.89 and 1.98; apices with
short spines or acute teeth, outer angles
sharply defined, right or slightly acute,
sutural angles obtuse; striae impressed,
scarcely punctulate; intervals convex,
slightly sparsely punctulate, 3rd usually 3-
punctate (4-punctate on left side only in in-
dividual from above Bernhard Camp).
Claw.') with 5 or 6 teeth. Secondary sexual
characters: i tarsi as genus; i middle
tibiae bent-in at apex but not tuberculate-
serrate; 6 with .3 or 4, 9 4 or 5 apical ventral
setae each side. Measurements: length 8.2-
9.2; width 2.6-3.0 mm.
Types, [lolotvpe 6 (U.S.N.M.) from
Iloliandia, West N. G. (J. W. Bongberg);
and paratypes as follows. West N. G.: 1
9 , Mountain slope above Bernhard Camp,
100 m, Apr. 19.39 (Toxopeus). N-E. N. (;.:
1 9 , Wan, Morobe Dist., 1200 m, Oct. 29,
1961 (Sedlacek); 1$, Erima, Astrolabe
Bay, 1896 (Biro). Pa|Mia: 1, Daradae,
near Javarere, Musgrove R., 100 m, Oct. 4,
1958 (Gressitt).
Additional material. N-E. N. G.: 1 9,
Tsenga, Upper Jimmi Vallev, 1200 m, Julv
14, 1955 (Gressitt).
Measured specimens. The i holotype and
the 9 paratype from Wau.
Notes. The acutely dentate or short-
spined rather than obtusely angulate elytral
apices distinguish this from the preceding
species (tripuncta). More material, espe-
cially a good series taken at one time and
place, is needed to show whether the dif-
ference is in fact specific.
Demefrida lafongula n. sp.
Description. With characters of genus;
form (Fig. 102) small and moderately broad
(in genus); reddish brown; not pubescent,
reticulate microsculpture present (some-
times faint, and slightly transverse ) only
on elytra, surface not much punctulate.
Head 0.89 and 0.91 width prothorax; eyes
prominent, genae short, not prominent;
front slightly irregularly impressed or with
punctiform impression before middle. Pro-
thorax subcordate, wide; width length 1.35
and 1.35; base apex 1..38 and 1.39; base'
head 0.97 and 0.94; sides broadly rounded
anteriorly, sinuate before well defined right
(sometimes slightly obtuse or acute) basal
angles; margins moderately wide, each with
seta before middle but none at base; disc
sometimes slightly wrinkled or subpunctate
basally and laterally. Elytra rather short
and broad (in genus); width elytra pro-
thorax 1.85 and 1.78; apices obtusely an-
gulate, with outer angles obtuse and usually
slightly blunted, sutural angles blunted;
striae imjiresscHl, \ aguely or not punctulate;
inter\'als convex, si)arsel\- pimctulate, 3rd
with 2 dorsal punctures. Clans with 3 or
4 teeth (and sometimes a small 5th one).
Secondary sexucd characters: i tarsi as
genus; ,5 middle tibiae strongK' tubercu-
late-serratc>; .s w ith 2 or 3, 9 3 or 4 apical
\ciitral setae each side. Measurements:
length 5.5-7.1; width 2.3-2.9 mm.
Types. Ilolotype $ (Bishop Mus.) from
Bisianumu, Iv of Port Mor(\sbv, Pajma,
500 m, Sejit. 3, 1959 (T. C. Maa); and
The Carabid Beetles of New Guinea • Darlington
159
paratypes as follows. Papua: 1 ? , Brown
R., E. of Port Moresby, 100 m, June 8,
1955 (Gressitt, now in M.C.Z., Type No.
.31,453); 1 9, Brown R., May 21, 1956
(E. J. Ford, Jr., Bishop Mus.); 1, Mt.
Lamington, 1300-1500 ft. (c. 400-460 m)
(C. T. McNamara, S. Australian Mus.); 1,
Buna Bay (C. T. McNamara, S. Australian
Mus.). N-E. N. G.: 1 9, Huon Pen.,
Pindiu, Apr. 20, 1963 (Sedlacek). West
N. G.: 1 c^ , Maffin Bay, Sept. 1944 (E. S.
Ross, Cal. Acad.).
Measured specimens. The i holotype and
1 9 paratype from Brown R.
Notes. This apparently widely distributed
lowland species is characterized by small
size, relatively broad form, and obtuse an-
gulation of elytral apices. See preceding
Key to Species of Demetrida for further
differential characters.
Demetrida angulata n. sp.
Description. With characters of genus;
form slender-average, with obtusely angu-
late elytral apices; reddish browai; not pubes-
cent, reticulate microsculpture distinct (light
and usually slightly transverse) only on
elytra, surface not much punctulate. Head
1.07 and 1.08 width prothorax; eyes promi-
nent, genae short, oblique, not prominent.
Prothorax subquadrate, narrow; width/
length 1.01 and 1.04; base/apex 1.40 and
1.31; base/head 0.88 and 0.87; sides weakly
irregularly arcuate (sometimes almost par-
allel) in c. anterior %, broadly sinuate be-
fore ± right but blunted posterior angles;
margins moderate, each with seta-bearing
puncture before middle but none at base;
disc variably wrinkled or subpunctate pos-
teriorly and laterally. Elytra: width elytra
prothorax 1.88 and 2.08; apices obtusely
angulate, outer angles sharply defined but
varying from slightly obtuse to acute,
sutural angles blunted; striae impressed,
usually faintly punctulate; intervals convex,
sparsely inconspicuously punctulate, 3rd
with 2 dorsal punctures. Claws with 5 or
6 teeth. Secondary sexual characters: S
tarsi as genus; i middle tibiae scarcely
modified, slightly bent-in near apex, not
distinctly tuberculate-serrate; S with 3 or 4,
9 5 or 6 setae each side last ventral
segment. Measurements: length 7.5-8.9;
width 2.5-3.1 mm.
Types. Holotype 6 (Bishop Mus.) from
Brown R., Papua, Sept. 30, 1959 (T. C.
Maa), taken sweeping; and paratypes as
follows (some in M.C.Z., Type No. 31,454).
Papua: 1 9 , same data as type except
dated Aug. 30, 1959; 1 9 , Brown R., E. of
Port Moresby, 100 m, June 8, 1955 (Gres-
sitt); 1, same locality, Apr. 27, 1960 (C.
W. Obrien, Bishop Mus.); 1 9, Laloki, nr.
Port Moresby, Aug. 30-Sept. 2, 1959 (T. C.
Maa, Bishop Mus.); 1 9, Kiunga, Fly R.,
July 11-14, 1957 (W. W. Brandt, Bishop
Mus.); 1 9, Daradae, nr. Javarere, Mus-
grove R., 100 m, Oct. 2, 1958 (Gressitt).
Measured specimens. The i holotype and
1st 9 paratype from Brown River.
Notes. See Key to Species of Demetrida
of New Guinea for distinguishing charac-
ters of an<i,ulata. It is the only New Guin-
ean Demetrida known to occur also in
Australia ( 1 9 , Rocky R., mid-peninsular
Cape York). In New Guinea, it has been
found only in Papua.
Demetrida reversa n. sp.
Description. With characters of genus;
form of preceding species (angulata) but
slightly less narrow; reddish brown; not
pubescent, reticulate microsculpture distinct
(and somewhat transverse) only on elytra,
but much of surface sparsely punctulate.
Head 1.03 and 1.07 width prothorax; eyes
prominent, genae short and oblique, not
prominent. Prothorax subquadrate; width/
length 1.10 and 1.12; base apex 1.32 and
1.31; base head 0.94 and 0.98; sides nearly
straight and subparallel or slightly con-
verging anteriorly, subangulate at setae,
broadly sinuate before right or slightly
acute basal angles; margins moderate, each
with seta-bearing puncture at or before
middle but none at base; disc slightly ir-
regularly subpunctate at base and laterally.
Elytra: width elytra/prothorax 1.85 and
160
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
1.83; apices ungulate (the angles c. right,
but variable), outer angles right or acute,
sharply formed; sutural angles obtuse-
blunted; striae impressed, faintly punc-
tulate; intervals sparsely punctulate, 3rd
with 2 dorsal punctures. Claws with 6 or 7
teeth. Secondary .sexual chaiacter.s: S tarsi
as genus; i middle tibiae slightly bent out
near apex, inconspicuously or irregularly
tuberculate-serrate (Fig. 162); i with c.
3, 9 c. 6 setae each side last ventral seg-
ment. Measurements: length 8.5-9.2; width
3.0-3.4 mm.
Types. Holotype 6 ( Bishop Mus. ) and
7 paratypes (3 in M.C.Z., Type No. 31,455)
from Guega, W. of Swart Valley, West
N. G., 1200 m, Nov. 14, 15, 1958 (Gressitt),
and 1 paratype, Swart Valley, W. Fork,
1300-1350 m, Nov. 17, 1958 (Gressitt).
Measured specimens. The 6 holotype and
1 9 paratype from Guega.
Notes. Among similar species with angu-
late but not spined elytral apices, this is
distinguished by quadrate prothorax, pro-
portions as given, and especially by form
of 6 middle tibiae, slightly bent outaard
at apex. Nevertheless the present species
may be closely related to the preceding one
[an^ulata), which is known only from
Papua (and Australia), while the present
one is known only from a restricted area
of West New Guinea.
Demefrida kokoda n. sp.
Descri])(ion. With characters of genus;
form as in Figure 103, large, slender;
reddish brown; not pubescent, reticulate
microsculpture present (light or iaint) onK-
on elytra, surface not much |)unctulate.
Head 0.99 and 0.99 width prothorax; exes
slightK' smaller than usual but prominent,
genae scarcely distinct from neck. Prothorax
cordate-subquadrate; widtli knigth 1.11 and
1.12; base/apex 1.22 and 1.23; base head
0.84 and 0.87; sides strongh' rounded in
anterior %, strongly sinuate before c. right
or slightly acute but blunted [)()st('rior
angles; margins narrow, each with seta
slightly before middle but none at base;
disc with middle line finer than usual in
genus, baso-lateral areas slightly punctate.
Elytra very long; width elytra ^ prothorax
1.72 and c. 1.86 (elytra spread); apices
with moderate spines, outer angles c. right
or slightly acute, sharply formed, sutural
angles slightly obtuse, sometimes denticu-
late; striae impressed, punctulate; intervals
only slightly convex, scarcely punctulate,
3rd with 2 seta-bearing punctures. Claws
with 7 or 8 teeth. Secondary .sexual char-
acters: 6 tarsi as genus; i middle tibiae
weakly tuberculate-serrate; i with c. 4, 9
numerous (up to 9) apical ventral setae
each side. Measurements: length c. 10.0-
11.0; width 2.9-3.5 mm.
Types. Holotype i ( British Mus. ) and
5 paratypes (2 in M.C.Z., Type No. 31,456)
from Kokoda, Papua, 1200 ft. (366 m).
May, Aug. (holotype), Sept., Oct., 1933
( Cheesman ) ; 1 paratype, Popondetta,
Papua, 25 m, June 1966 ( Shanahan-Lip-
pert. Bishop Mus.).
Additional material. N-E. N. G.: 1 9,
Wau, Morobe Dist., 1050 m, Apr. 30, 1962
(Sedlacek). West N. G.: 1 9, Waris. S.
of Ilollandia, 450-500 m, Aug. 24-31, 1959
(T. G. Maa, Bishop Mus.); 1 9 , Hollandia.
J:m. 1945 (B. Malkin, U.S.N.M.).
Measured sj)ecimens. The c5 holotype and
1 9 paratype.
Notes. The form ol this species is uni({ue
in the genus, so far as I know, and other
characters including the relatixely fine
middle line ol the pronotum are distinctixe.
Demetrida moda n. sp.
Description. See Phite 1, figure I\'; with
characters of genus; reddish brown; not
pubesci'ut, reticulate microsculpture distinct
(but light) onl\' on elytra, parts ol upper
siu"lace sparsely j)unctulate. Head 0.85
and 0.84 width prothorax; e\'es prominent,
genae short, obliciue. not jirominent. Vro-
thorax subcordalc; width length 1.46 and
1.33; base /apex 1.40 and 1.37; base/head
1.02 and 1.04; sides arcuate^ anteriorly,
strongly sinuate belorc sharply defined right
or acute posteiioi- angles; margins moderate.
The Carabid Beetles of New Guinea • Darlington
161
each with seta before middle but none at
base; disc slightly punctate basally. Elytra:
width elytra prothorax 1.69 and 1.77; apices
with moderate spines, outer angles obtuse,
sutinal angles obtuse; striae impressed,
faintly punctulate; 3rd intervals 2-punctate.
Claws with c. 4 teeth. Secondary sexual
characters: 6 tarsi as genus; 6 middle
tibiae strongly tuberculate-serrate ( c. 4
rounded tubercles, Fig. 160); i with 2, 9
3 apical ventral setae each side (number
may vary). Measurements: length c. 5.5-
6.5; width 2.1-2.5 mm.
Types. Holotype S (M.C.Z., Type No.
31,457) and 4 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington), and
additional paratvpes as follows. Papua: 8,
Kokoda, 1200 ft.' (366 m), Aug., Sept., Oct.,
1933 (Cheesman). N-E. N. G.: 2, Sattel-
berg, Huon Gulf, 1899 (Biro).
Additional material N-E. N. G.: 2,
Pindiu, Huon Pen. (1 labeled 500-600 m),
Apr. 19, 20, 1963 (Sedlacek); 1, Finschhafen,
May 7, 1944 (E. S. Ross, Gal. Acad.); 1,
Bubia, Sept. 1949 ( N. L. H. Krauss, Bishop
Mus.).
Measured specimens. The 6 holotype and
1 9 paratype from Dobodura.
Notes. This species, as the name moda
is intended to suggest, is the first of several
generally similar forms which differ among
themselves slightly in proportions and color
and more significantly in length of elytral
spines and punctation of 3rd elytral inter-
vals. Some of these forms may be geo-
graphically limited and allopatric and may
eventually be considered subspecies. The
present species seems to be confined to the
eastern half of New Guinea. Its differential
characters are given in the preceding Key
to species.
Demefrida submoda n. sp.
Description. With characters of genus;
form of preceding species (rnoda) except
elytra acutely toothed, not spined, and
proportions slightly different, with head rel-
atively slightly wider and base of prothorax
narrower; color, microsculpture, etc. as in
moda. Head 0.89 and 0.91 width prothorax;
eyes prominent, genae short and oblicjue.
Prothorax subcordate; width/length 1.40
and 1.37; base/apex 1.43 and 1.45; base/
head 0.97 and 0.95; sides rounded anteriorly,
strongly sinuate before c. right posterior
angles; margins rather wide, each with seta
near or before middle but none at base;
disc slightly irregular or subpunctate baso-
laterally. Elytra: width elytra prothorax
1.70 and 1.81; apices acutely angulate or
denticulate, outer angles obtuse or blunted,
sutural angles obtuse; striae impressed, not
distinctly punctulate; intervals convex,
slightly inconspicuously punctulate, 3rd 2-
punctate. Claws with c. 5 teeth. Secondary
sexual characters: 6 tarsi as genus; 6 mid-
dle tibiae tuberculate-serrate (4 tubercles);
6 with 2, 9 3 apical ventral setae each
side. Measurements: length 6.0-7.0; width
2.3-2.8 mm.
Types. Holotype S (Bishop Mus.) and
17 paratypes (7 in M.G.Z., Type No.
31,458) from Wau and vicinity (including
Mt. Missim), Morobe Dist., N-E. N. G.,
altitudes from 950 to 1400 m, dates in Jan.,
Feb., Mav, Julv, Aug., Sept., Nov., Dec,
1961-1964 (Sedlaceks, T. G. Maa) (holo-
type, 1250 m. May 3, 1963); and additional
paratvpes from N-E. N. G. as follows: 2,
Bulolo, 1065 m, Aug. 15, 16, 1956 (E. J.
Ford, Jr., Bishop Mus.); 1, Upper Watut
R., 24 km W. Bulolo, 760 m. Mar. 5-6,
1963 (Sedlacek).
Additional material. N-E. N. G.: 1, vie.
Nadzab, July 1944 (Darlington); 2, Kas-
sem, 48 km E. of Kainantu, 1350 m, Nov.
7, 1959 (T. G. Maa, Bishop Mus.); 1,
Kumun, Upper Jimmi Valley, 1000 m, July
13, _ (Gressitt); 1, Maprik, 150 m, Dec.
29-Jan. 17, 1960 (T. G. Maa, Bishop Mus.);
1, Eliptamin ^^^, 1200-1350 m, July 16-31,
1959 (W. W.' Brandt, Bishop Mus.); 1,
Goroka, 1550 m, June 19, 1955 (Gressitt),
"pigeon peas cane." West N. G.: 1, Hol-
landia, Dec. 15, 1944 ( Hoogstraal, M.G.Z.).
Measured s})ecinwns. The 6 holotype and
1 9 paratype from Wau.
Notes. This species differs from moda
162 Bulletin Museum of Coinparative Zoology, Vol. 137, No. 1
as indicated in tht' preceding Description.
It may prove to be only a subspecies of
moda, and has thus far been found only in
the central-eastern part of the north side
of New Guinea, chieflv in the lower moun-
tains.
Demetrida hollandia n. sp.
Description. With characters of genus;
form c. of moda and suhmoda (above) but
color darker, reddish brown with elytra
darker brown or brownish black with apical
% or less often paler, the pale apical area
varying in distinctness and extent; micro-
sculpture, etc. c. as in moda, with reticula-
tions faint but usually visible on elytra.
Head 0.89 and 0.(S8 width prothorax, eyes
prominent, genae short. Prothorax sub-
cordate, slightly narrower than in moda;
width length 1.32 and 1.31; base/apex 1.36
and 1.34; base head 0.97 and 0.95; sides
slightly irregularly rounded, often subangu-
late at lateral setae; margins moderately
wide, each with seta near or before middle
but none at base; siuface not or ver\ Httle
punctate. Elytra: width elytra prothorax
1.86 and 1.85; apices short-spined, outer
angles obtuse but more distinct than in
moda, sutural angles blunted; striae im-
pressed, not distinctly punctulate; intervals
convex, scarcely punctulate, 3rd with 2
principal punctures and often (not alwa\s)
with 1 or more intermediate i^unctures
which vary in size and sometimes do and
sometimes do not bear setae. CUaus with
c. 4 te(>th. Secondary sexital characters: i
tarsi as genus; i middle tibiae strongly
tuberculate-serrate (c. 4 tubercles); i with
2, 9 3 apical ventral setae each side. Mea-
surements: length 5.8-7.6; width 2.1-2.9
mm.
Types. Tlolotype S (M.C.Z., Type No.
31,459) and 33 paratypes from vie. ITol-
laudia. West N. (i., July-Sept. 1944 (Dar-
lington ).
Addilional material. West N. (',.: 11,
Hollandia and vicinity including C]yclops
Mts., at low altitudes (not over 500 m),
\ari()us dates and collectors; 10. Maffin
Bav, dates in June, July, Aug., Sept., Oct.
1944 (E. S. Ross, Cal. Acad.); 5, Nabire,
S. Geelvink Bay, 5-50 m, Aug. 25-Sept. 5.
1962 (Sedlacek); 1, \\'a.sian (Vogelkop),
Sept. 1939 (Wind, M.C.Z.); 1, Fac Fac,
June 1939 (Wind, M.C.Z.). N-E. N. G.:
33, various localities including Fluon Pen.;
Torricelli Mts.; Sepik Dist.; Wewak; Lae;
Bulolo; Wau. Papua: 1 teneral 6 , doubt-
fully identified, from Bisianumu, E. of Port
Moresby, 500 m, Sept. 24, 1955 (Gressitt).
Measured specimens. The c5 holotype and
1 9 paratype.
Notes. D. hollandia seems closely related
to moda and suhmoda but is distinguished
by characters given above. Some of the
specimens listed under Additional material
\ar\- toward icaii (see below).
Demetrida wau n. sp.
Description. With characters of genus;
form c. as in moda, sidmwda, and hollandia
but more slender with eyes less prominent
and genae slightly longer and less abrupt
than in the species named; reddish brown,
eU'tra brownish black, usualK' not paler
at apex; not pubescent, upper surface in-
cluding elytral disc without reticulate micro-
sculpture but in part (especialK- cKtra)
sparseK' punctulate. Head 0.89 and 0.89
width prothorax; e>'es and genae as indi-
cated abo\e. Prothorax subcordate; width/
length 1.21 and 1.22; base apex 1.44 and
1.44; base head 0.99 and 0.99; sides rounded
anteriorly, often subangulate at setae (as in
hollandia), strongly sinuate before c. right
basal angles; margins rath(M- wid(\ each
with seta near or beloic middle but none
at base; disc more punctate basalh' and
laterally than in the '^^ preceding species.
Elytra: width elytra prothorax 1.7.3 and
1.73; a|Mees short-spined (or with long
acute teeth), with outer angles obtuse
but distinct, sutural angles blunted; striae
impressed, punctulate; interxals eou\('x,
punctulate. 3i(l olten 4-i')unetate but inter-
mediate |iuiutui('s \aiiabl(' in si/e and
sometimes absent and with oi' without
setae. Clans with c. ,5 teeth. Secondary
The Carabid Beetles of New Guinea • Darlingtofi
163
sextial cJiamcters: 6 tarsi as genus; 6
middle tibiae weakly tuberculate-serrate (c.
3 tubercles distinct); 6 with 2, 9 3 apical
ventral setae each side. Measurements:
length 6.4-7.9; width 2.4-2.9 mrn.
Types. Holotype c^ (Bishop Mus.) and
118 paratypes (some in M.C.Z., Type No.
31,460) all from Wau, Morobe Dist., N-E.
N. G.; altitudes from 1000 to 1450 m;
dates in everv month, 1961-1963 (holotvpe,
1200 m, July '22, 1961 ) ( Sedlaceks ) .
Additional material. N-E. N. G. : 9 addi-
tional teneral, broken, or atypical speci-
mens from Wau; 1, Jim(m)i R., E. High-
lands, Julv-Sept. 1961 (W. W. Brandt,
C.S.I.R.O.); 1, Upper Watut R., 24 km W.
Bulolo, 760 m, Mar. 5-6, 1963 (Sedlacek);
1, Erima, Astrolabe Bay, 1897 (Biro). West
N. G.: 2, Hollandia, May, June 1945 (B.
Malkin, U.S.N.M.); 1, Waris, S. of Hol-
landia, 450-500 m, Aug. 16-23, 1959 (T. C.
Maa, Bishop Mus.); 1, Ifar, 400-550 m,
June 23, 1959 (T. C. Maa, Bishop Mus.).
Measured specimens. The S holotype and
1 9 paratype.
Notes. D. wau may be primarily a geo-
graphic representative of liollandia, but the
long type series seems distinct; the speci-
mens listed above from Hollandia and
Waris are plainly wau, not hollandia; and
the one from Ifar seems to be wau except
that the elytra are distinctly microreticulate.
Apparent intermediates do occur at some
other localities, however. They are tenta-
tively placed with Additional material un-
der hollandia. See also Notes under D.
subpunctata (3rd species below).
One 9 of wau, from Wau, is a note-
worthy abnormalit)', \\4th the posterior pro-
thoracic angles irregularly widened and
each \\'ith 2 setae, although normal in-
dividuals of wau lack posterior-lateral
setae.
Demefrida similis n. sp.
Description. With characters of genus;
form of moda, etc. but larger, rather slender,
with prothorax narrowly subcorclate and
elytra spined or acutely dentate and with
outer angles sharply formed; reddish
brown, elytra not or only slightly darker;
not pubescent; reticulate microsculpture
visible (often faint) only on elytra. Head
0.87 and 0.90 width prothorax; eyes promi-
nent, genae shorter, oblique. Prothorax
subcordate; width/length 1.26 and 1.22;
base/apex 1.35 and 1.34; base/head 0.98
and 0.98; sides broadly sometimes slightly
irregularly arcuate in more than % of length,
strongly sinuate before right or slightly
acute usually slightly blunted posterior
angles; margins rather wide, each with
seta-bearing puncture before middle but
none at base; disc slightly punctate at
base and sides. Elytra long; width elytra/
prothorax 1.61 and 1.69; apices short-
spined (rarely only acutely toothed), outer
angles well formed, varying from slightly
obtuse to acute, sutural angles blunted-
obtuse; striae impressed, finely punctulate;
intervals slightly convex, sparsely finely
punctulate, 3rd 2-punctate ( all specimens ) .
Claws with 6 or 7 teeth. Secondary sexual
characters: 6 tarsi as genus; S middle
tibiae tuberculate-serrate (c. 4 tubercles);
S with 2, 9 3 or 4 apical ventral setae each
side. Measurements: length 8.8-10.8; width
3.0-3.5 mm.
Types. Holotype £ (M.C.Z., Type No.
31,461 ) and 3 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratvpes as follows. Papua: 4,
Kokoda, 1200, 1300 ft. {c. 366, 400 m),
June, Aug., Sept. 1933 (Cheesman), 1
labeled also 'Tn fungus. A, & under bark
behind it," and 1 "At light"; 1, same locality,
380 m. Mar. 20, 1956 (Gressitt), in light
trap; 2, Kokoda-Pitoki, 400 m. Nhu". 23,
1956 (Gressitt); 2, Mt. Lamington, 1300-
1500 ft. (c. 400-460 m), (C. T. McNamara,
S. Australian Mus.).
Additional material. Papua: 1, Kiunga,
Fly R., Oct. 1-7, 1957 (^^^ ^^^ Brandt,
Bishop Mus.). N-E. N. G.: 1, Ebabaang,
Mongi W\itershed, Huon Pen., 1300-1400
m, Apr. 16-18, 1955 ( E. O. Wilson, M.C.Z.).
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
164 Bulletin Museum of Couiparativc Zoology, Vol. 137, No. 1
Notes. Characters distinguishing this
species from moda, etc. are given in the
Description, above; and see also Notes
under the following species.
Demefrida duplicafa n. sp.
Description. With characters of genus;
form c. as in similis (above); reddish
brown, elytra not or not much darker; not
pubescent, reticulate microsculpture visible
only on elytra, more transverse than in
similis, surface in part sparsely punctulate.
Head 0.92 and 0.91 width prothorax; eyes
prominent, genae shorter, oblique. Pro-
tJiorax quadrate-subcordate; width length
1.27 and 1.29; base/apex 1.31 and 1.31;
base/head 0.94 and 0.96; sides (usually a
little irregularly) rounded anteriorly, sinu-
ate before c. right but usually blunted pos-
terior angles; margins rather wide, each
with seta at or slightly before middle but
none at base; disc not much punctate
even basally. Elytra: width elytra/ pro-
thorax 1.71 and 1.75; apices with moder-
ate spines, outer angles sharply defined
and sometimes acutely denticulate, sutural
angles blunted-obtuse; striae moderately
impressed, scarcely punctulate; intervals
slightly convex, 3rd with 2 principal and
usually one or more smaller intermediate
dorsal punctures. Claws with c. 5 teeth.
Secondary sextial characters: i tarsi as
genus; 6 middle tibiae tubereulate-serrate
{c. 6 small tubercles); 6 with 2, 9 3
apical ventral setae each side. Measiire-
incnts: length c. 8.0-9.0; width 2.9-3.3
mm.
ry))es. llolotype $ (M.C.Z., Type No.
31,462) and 10 paratypes from Dobodiua,
l^apiia, Mar.-July 1944 ( Darhiiglon ); 3
[)aratyp('s bom Kokoda, Papua, 1200 It.
(366 m), Apr., June, Aug. 1933 (Cheesman);
94 paratypes, Mt. Lamington, Papua, 1300-
1500 ft. {c. 400-460 m) (C. T. McNamara,
S. Australian \bis. ).
AddHio)ud nialcriid. Si.\t)-one (includ-
ing 44 from W'au), from 12 localities, in all
3 political divisions of New (guinea ( from
Milne lia\ to mountain slojX' al)o\c l^crn-
hard Camp), altitudes from near sea level
to at least 1500 m (at Wau), various dates
and collectors.
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. D. duplicata is much like similis
and occurs at some of the same localities
l)ut differs constantly (at least at Dobodura)
by having the elytra more distinctly and
more transversely microreticulate, with 3rd
intervals with more than 2 dorsal punctures.
The additional punctures vary in size and
sometimes do and sometimes do not bear
setae.
At Dobodura, duplicata (like similis) is
uniformly brown, but individuals with base
of elytra ± darker occur with brown in-
di\ iduals at many localities including Wau.
The color is not obviously dimorphic but
apparently continuously variable. A re-
lated population in \\hich the elytra are
always dark at base occurs in West N. G.
(see hasalis, p. 172). The specimens sum-
marized above under Additional material
vary in other ways which cannot profitably
be discussed in detail here.
Demefrida subpuncfafa n. sp.
Description. ^Vith characters of genus;
fomi c. as in moda, udti. etc., but slightly
more slender; dark reddish brown, elytra
darker (dark castaneous), legs browii; not
pubescent; microsculpture \isible ( faint,
distinctK' trans\cM"se ) ouK on eUtra, but
much of upper surface finely sparsely
punctulate. Head 1.00 and 0.96 width i^ro-
thorax; eyes moderate, genae slightb'
shorter, obliciue. Prothorax narrowly sub-
cordate; width length 1.16 and 1.16; base
apex 1.37 and 1.3(S; base head 0.93 and
().9(S; sides weakly irregularly arcuate in
anterior "i or more, strongK' sinuate before
right or slightly acute i)()sterior angles;
margins narrower than in moda and irau.
each with seta before middle but none at
base; sinface rather closely punctate across
base and in margins. Elytra: width elytra
prothorax 1.8(S and I.SS; ai:)ices with short
spines, outer angles distinct but obtuse ami
The Carabid Beetles of New Guinea • Darlington 165
sometimes slightly blunted, sutural angles
blunted or narrowly rounded; striae im-
pressed, faintly punctulate; intervals con-
vex, 3rd usually 3- ( rarely 4- ) punctate but
intermediate puncture(s) variable in size
and sometimes indistinguishable. Claws
with c. 4 teeth. Sccondanj sexual characters:
i tarsi as genus; i middle tibiae tubercu-
late-serrate ( c. 4 rounded tubercles ) ; S
with 2, 9 3 apical ventral setae each side.
Measurements: length 6.7-7.6; width 2.3-
2.7 mm.
Types. Holotype S (M.C.Z., Type No.
31,463) and 5 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes from Papua as fol-
lows: 7, Kokoda, 1200 ft. (366 m), May,
June, Julv, Aug., Oct. 1933 (Cheesman);
5, Kokoda-Pitoki, 450 m. Mar. 24, 1956
(Gressitt); 1, Bisianumu, E. of Pt. Moresby,
500 m, Sept. 23, 1955 (Gressitt); 1, "Papua,"
(Hungarian Nat. Mus. ); 3, Mt. Lamington,
1300-1500 ft. (c. 400-460 m) (G. T. Mc-
Namara, S. Australian Mus.).
Measured specimens. The S holotype and
1 9 paratype from Dobodura.
Notes. This may ( or may not ) be the
Papuan representative of the hollandia-icau
group of central and western New Guinea.
D. suhpunctata most resembles wau but is
slightly more slender, with narrower pro-
thoracic margins ( which distinguish it also
from hollandia), and with distinct elytral
microsculpture.
Demetrida dobodura n. sp.
Description. With characters of genus;
form c. as in moda and similis but slightly
more slender; brown (not dark), elytra not
or not much darker; not pubescent, reticu-
late microsculpture visible ( light or faint )
only on elytra, surface (except of elytra)
not much punctulate. Head 0.98 and 0.96
width prothorax; eyes prominent, genae
short, oblique. Prothorax subquadrate;
width/length 1.11 and 1.12; base/apex 1.34
and 1.30; base/head 0.95 and 0.96; sides
weakly irregularly arcuate in % or more of
length, weakly sinuate before c. right but
blunted posterior angles; margins narrower
than in moda and .similis, each with seta at
or slightly before middle but none at base;
surface weakly punctate across base and in
margins. Elytra: width elytra/prothorax
l.ScS and 1.87; apices with moderate spines,
outer angles acutely denticulate (or right
but sharply formed in some individuals
listed under Additional material), sutural
angles blunted-obtuse; striae moderately im-
pressed, finely punctulate; intervals slightly
convex, punctulate, 3rd with 2 dorsal punc-
tures. Cdans with c. 5 teeth. Secondary
sexual characters: i tarsi as genus; i
middle tibiae tuberculate-serrate (3 or 4
\\'ell spaced small tubercles ) ; S with 2 or
3, 9 5 to 7 apical ventral setae each side.
Measurements: length 7.7-9.2; width 2.6-
3.3 mm.
Types. Holotype S (M.C.Z., Type No.
31,464) and 21 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
9 paratypes, Kokoda, Papua, 1200 ft. (366
m), Aug., Sept. 1933 (Gheesman).
Additional material. Papua: 1, Kiunga,
Fly R., Aug. 8-10, 1957 (^^^ \\'. Brandt,
Bishop Mus.); 1, Koitaki, 1500 ft. (c. 450
m), Oct.-Nov. 1928 (Pemberton, H.S.P.A.).
N-E. N. G.: 1, Pindiu, Huon Pen., 500-600
m, Apr. 19, 1963 (Sedlaeek). Also 1 old
specimen, i , labeled "New Guinea. Sayer,"
"probably N. gen. near Euproctus," and
"Gen. probably near Gtenodactylus" (the
last 2 labels probably by Andrewes ) .
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. Among other brown, spined
Demetrida. this should be recognizable by
prothorax subquadrate c. wide as head and
slightly wider than long by measurement,
by the rather large size, and the 2-punctate
3rd elytral intervals. See also Notes under
following species (kiunga).
The specimens listed under Additional
material are doubtfully identified. D.
dobodura is therefore known with certainty
only from Dobodura and Kokoda, in
Papua.
166 BiilJctin Mu-scmn of Companitive Zoology, Vol. 137, No. 1
Demefrida kiunga n. sp.
Description. With characters of genus;
form c. of preceding (dohocJum) but
larger; reddish brown, elytra slightly but
not much darker; not pubescent, reticulate
microsculpture distinct only on elytra, sur-
face not much punctulate. Head 1.04
and 1.00 \\idth prothorax; eyes moderately
prominent, genae shorter and oblique.
Pmtliorax subquadrate; width length 1.00
and 1.02; base/apex 1.31 and 1.24; base/
head 0.90 and 0.89; sides very weakly
arcuate anteriorly, slightly subangulate at
setae, strongly sinuate well before slightly
acute sometimes slightly blunted basal
angles; margins rather narrow, each with
seta at or before middle but none at base;
disc slightly irregular or subpunctate at base
and in margins. Elytra long; width elytra/
prothorax 1.87 and 1.80; apices long-spined,
outer angles c. right and sharply formed
but not denticulate, sutural angles obtuse;
striae impressed, faintly punctulate; inter-
\als slightly convex, 3rd 2-punctate. Claws
with c. 7 teeth. Secondanj sexual ehar-
aeters: S tarsi as genus; c^ middle tibiae
unmodified, virtually straight, not tubercu-
late-serrate; i with 3 or 4,9 4 or 5 ( un-
symmetric in both individuals ) setae each
side last ventral segment. Measurements:
lengdi 10.8; width 3.4 mm.
Tijpes. Holotype <^ (Bishop Mus.) and
1 9'paratype (M.C.Z., Type No. 31,465)
from Kiunga, Fly R., Pajma, Aug. 14-17,
18-23, 1957 (W. W. Brandt).
Notes. Among other jilain reddish brown
Demetrida with (piadrate prothorax and
spined elytra, this is distinguished by rel-
atively large si/e, proportions, simple $
middle tibiae, and oilier characters given
in the Key to Species.
As compared with dohodtira, the present
species is larg(>r, with longer elytral spines
but less produc{>d outer elytral angles, as
well as with differcMit ^ tibiae. The single
individual of dobodura seen from Kiunga
is a i with all the characters of dobodura:
smaller si/.e, denticulate outer elytral angles.
and plainly tubcrculate-serrate middle
tibiae.
Demefrida mafuiu n. sp.
Description. See Plate 2, figure V; with
characters of genus; color dimorphic, either
dark red with prothorax and basal -.-. of
elytra green with green color extending
farther back at sides than at suture or
entirely irregular dark reddish brown, legs
either dark with paler tarsi or entirely
brown, antennae brown in both cases; not
pubescent, reticulate microsculpture pres-
ent, c. isodiametric on head and elytra and
transverse on pronotum, surface not much
punctulate. Head 1.11 and 1.07 width pro-
thorax; eyes prominent, genae shorter,
oblique, not prominent; front flattened and
irregularly slightly impressed before mid-
dle. Prothorax subquadrate; width length
0.96 and 0.99; base apex 1.35 and 1.35;
base/head 0.84 and 0.87; sides weakly
arcuate for much of length, scarcely angu-
late at setae, sinuate before prominent c.
right basal angles; margins narrow, each
with seta-bearing puncture slighd>' before
middle but none at base; disc more convex
than usual, baso-lateral impressions irregu-
larly punctate. Elytra: width elytra/pro-
thorax 1.96 and 2.03; apices acutely angu-
late or dentate, outer angles sharply formed,
c. right or obtuse, sutural angles obtuse-
striae well impressed, fineU' punctulate;
intervals slighth' con\ex, faintl)' sparsely
punctulate, 3rd 2-punctate. Clatvs \\'ith c.
6 t(>eth. Secondary sexual characters: $
tarsi as genus; S middle tibiae hcut in at
apex but not tubcrculate-serrate; S with
4, 9 c. 5 apical ventral setae each side.
Measuremenis: length 9.3; width 2.9 mm.
Type. Ilolotvpc $ (British Mus.) from
Mafuiu, Papua, 4000 ft. (1220 m), Dec.
1933 (Cheesman); and 1 9 paratype (also
British Mus.) widi .same data v\cv\A datcnl
Ian. 1934.
Notes, (.'omparisoii with direr.sa (p.
172) suggests that the color dimorphism ot
nuifulu is simply MeiuU'lian. not sexual.
These 2 species max l)e related, but mafuUi
The Carabid Beetles of New Guinea • Darlington 167
seems surely distinct ])y form, greater
convexity of pronotum, and more distinct
reticulate microsculpture of much of the
upper surface. The 2 individuals of majuJu
share these characters and, except in color,
differ only slightly in other ways: e.g..,
the 9 has the elytral apices more acutely
toothed but the outer angles more obtuse.
The 2 color forms of mafulu are keyed out
separately in the Keij to Species.
Demetrido forma n. sp.
Description. With characters of genus;
form as in Figure 104; reddish brown,
legs testaceous; not pubescent, microsculp-
ture present ( weak or faint ) only on elytra,
surface not much punctulate. Head 1.12
and 1.10 width prothorax; eyes prominent,
genae short, oblique. Prothorox subquad-
rate, long; width length 0.99 and 1.01; base
apex 1.37 and 1.23; base/head 0.82 and
0.81; sides weakly irregularly rounded
through much of length, moderately sinu-
ate posteriorly before c. acute but blunted
basal angles; margins narrow, each with
seta-bearing puncture slightly before mid-
dle but none at base; surface in part ir-
regular or weakly punctate posteriorly and
laterally. Elytra rather long; width elytra/
prothorax 2.11 and 2.18; apices spined, outer
angles c. right, sharply formed, sutural
angles right or slightly obtuse, sometimes
slightly blunted; striae impressed, weakly
punctulate; intervals convex, 3rd with 2
dorsal punctures. Claws with 7 or 8 teeth.
Secondary sexual characters: S tarsi as
genus; 6 middle tibiae tuberculate-serrate
(c. 3 or more tubercles); 6 with 2 or 3, $ c.
4 apical ventral setae each side. Measure-
ments (type series): length 9.3-9.6; width
3.1-3.3 mm (specimens listed under Addi-
tional material 6.8-9.6 mm long).
Types. Holotype $ (Bishop Mus.) and
7 paratypes (3 in M.C.Z., Type No. 31,466)
all from Pindiu, Huon Pen., N-E. N. G.,
500-600, 870-1300 m; dates in Apr. 1963
(holotype, 500-600 m, Apr. 19) (Sedlacek).
Additional material. Twenty-three speci-
mens from 9 localities in Papua, N-E.
rV. G., and eastern West N. G. Because
of variations (see following Notes) and
doubtful identifications these specimens
are not recorded in detail.
Measured specimens. The i holotype and
1 9 paratype.
Notes. Under this species I have tenta-
tively placed all nonpubescent, brown,
pale-legged New Guinean Demetrida with
spined elytra, prothorax elongate-subquad-
rate and considerably narrower than head
( width head prothorax usually but not
always c. 1.10 or more), and 3rd intervals
2-punctate ( but see below ) . The specimens
thus assembled vary considerably in size,
prominence of eyes, exact form of prothorax,
and length of elytral spines. Several species
may be represented but, if so, I cannot
separate them now.
Although the 3rd intervals are 2-punctate
on both elytra in most individuals, in 3
cases a 3rd (intermediate) puncture is
present on one side only, and an individual
from Wau which I tentatively assign to
forma is 3-punctate on both sides. This
individual is the only forma (if it is this
species) seen from Wau. It is a small S
with tuberculate-serrate middle tibiae.
Demetrido recto n. sp.
Description. With characters of genus;
form c. as preceding species (forma) except
elytra short-spined; reddish brown, legs
pale with dark knees; not pubescent, reticu-
late microsculpture distinct only on elytra,
but surface in part finely sparsely punc-
tulate. Head 1.14 and 1.20 width prothorax;
eyes prominent, genae shorter, oblique, not
prominent (but see Notes below). Pro-
thorax long-quadrate; width length 0.92
and 0.88; base apex 1.28 and 1.28; base/
head 0.85 and 0.84; sides virtually straight
anteriorly or weakly angulate at setae,
sinuate well before slightly acute basal
angles; margins narrow, each with seta
slightly before middle but none at base;
baso-lateral impressions weak, subpunctate.
168 Bulletin Miiscuin of Comparative Zoology, Vol. 137, No. 1
Elytra: width elytra prothorax 2.14 and
2.30; apices shoit-.spined or acutely toothed,
outer angles sharply formed, acute, inner
angles obtuse; striae impressed, punctulate;
intervals convex, 3rd 2-punctate. Clows
with c. 6 or 7 teeth. Secondary .sexual
characters: 6 tarsi as genus; i middle
tibiae bent in at apex but not tuberculate-
serrate; 6 with 3, 9 4 or 5 setae each
side last ventral segment. Measurements:
length 8.4-9.7: width 2.7-3.3 mm.
Types. Holotype c^ (Bishop Mus.) and
2 9? paratypes (1 in M.C.Z., Type No.
31,467) from Wan, Morobe Dist, N-E.
N. G., 1200 (liolotypc), 1050, and 1090 m,
dates in Jan. 1963 (holotype, Jan. 8-10)
( Sedlacek ) .
Additional material. West N. G. : 1 c^ ,
Hollandia, May 1945 (Hoogstraal, M.C.Z.).
Measured sj^ccimens. The J, holotype and
1 9 paratype.
Notes. D. recta resembles forma but is
distinguished by straighter sides of pro-
thorax and especially by bent-in but not
tuberculate-serrate i middle tibiae. D.
recta may be more closely related to
l<iun£ia but is smaller, more slender, with
shorter elytral spines. The real interrela-
tionships of these and other more or less
similar species are doubtful.
Tliis species emphasizes that the ratio
base/apex of prothorax must be interpreted
with caution. The ratio of 1.28 in recta
suggests that the apex is considerably nar-
rower than the base, and this is true when
the apex is measured in the standard way,
between the most advanced points of the
angles. Nevertheless, the prothorax appears
\ iitually rectangular.
Th(> genae of the lK)l<)t\'|)e are unsym-
metrie: the right one is normal, as de-
scribed above and as in the other speci-
mens of the species, wlule the left one is
subangulate just behind the eye, although
not so jirominent as in tri})uncfa and
iS,enicula. This slight angulation of the left
gena in one specimen only ol recta is
presumably an abnormality.
Demetrida rex n. sp.
Description. With characters of genus;
form as in Figure 105; reddish brown; not
pubescent, reticulate microsculpture faint
or absent even on elytra, but much of
upper surface sparsely inconspicuously
punctulate. Head 1.06 and 1.08 width pro-
thorax (at middle); eyes prominent, genae
shorter, oblique. Prothorax trapezoidal,
wider at base than at middle; width ( at
middle) length 1.06 and 1.02; base, apex
1.41 and 1.34; base/width at middle 1.04
and 1.06; base head 0.98 and 0.98; sides
shaped as figured, narrowly margined, each
with seta-bearing puncture at or slightly
before middle but none at base; surface
irregularly slightly punctate at base and
sides. Elytra ample; width elytra prothorax
(at middle) 2.00 and 2.12; apices spined,
outer angles acutely denticulate, sutural
angles obtuse-blunted; striae impressed,
punctulate; intervals slightly convex, 3rd
with 3 dorsal punctures (all specimens).
Claws with 7 or 8 teeth. Secondary sexual
characters: 6 tarsi as genus; c5 middle
tibiae slightly bent-in toward apex but not
tuberculate-serrate; 6 with 2 or 3, 9 c. 5
apical ventral setae each side. Measure-
ments: length 10.2-11.4; width 3.5-4.0 mm.
Types. Holotype i ( Bishop Mus. ) from
Mokai Village, Torricelli Mts., N-E. N. G.,
750 m, Dec. 8-15, 1958 (^^^ W . Brandt);
and additional paratypes as follows. N-E.
N. G.: 1 9, Eliptaniin Vy., 1200-1350 m.
June 19-30, 1959 (W. \V. Brandt. Bishop
Mus.); 1 9, Adalbert Mts.. Wanunia, 800-
1000 m, Oct. 24, 1958 (Gressitt); 1 i , Pin-
diu, Huon Pen., Apr. 20, 1963 (Sedlacek).
Papua: 1 i , Dogon, Amazon Ba\ Dist..
2400 ft. ( c. 730 m), Oct.-NoN . 1962 (W. W.
Brandt, C.S.I.H.O.); 1 9, Owen Stanley
Rge., Ooilala, B()m(\ 1950 m. Apr. 16-30.
1958 (\\. W. Brandt. Bishop Mus.). (Thc>
]")aratvpes from Adalbert Mts. and Pindiu
now in M.C.Z.. Type No. 31,468.)
Measured specimens. The ■: holotype and
9 paratype from Eliptamin Valley.
Notes. Although the 6 speeiinens listi-d
The Carabid Beetles of New Guinea
Darlington
169
above come from 6 different localities, they
seem to be conspecific and to represent a
very distinct species, characterized by large
size, form of prothorax, presence of an
acute tooth ( almost a short spine ) at outer-
apical elytral angle, and 3-punctate 3rd
intervals. See Key to Species for place of
rex among other New Guinean Demetrida.
Demefrida brunnea n. sp.
Description. With characters of genus;
form average, with spined elytra; brownish
testaceous, head and prothorax usually
slightly darker than elytral disc, legs pale;
not pubescent, reticulate microsculpture
visible (light) only on elytra, surface not
much punctulate. Head 1.12 and 1.13 width
prothorax; eyes prominent, genae shorter,
oblique. Prothorax quadrate; width length
1.02 and 0.99; base/apex 1.23 and 1.26;
base/head 0.83 and 0.87; sides subparallel
or weakly irregularly arcuate in anterior %
or more, subangulate at lateral setae,
broadly sinuate before right or slightly
acute posterior angles; margins narrow,
each with seta at or slightly before middle
but none at base; surface irregular or
slightly punctate at base and sides. Elytra:
width elytra prothorax 2.06 and 2.14; apices
spined, outer angles denticulate, sutural
angles right or slightly obtuse, slightly
blunted; striae impressed, faintly punctu-
late; intervals slightly convex, 3rd usually
3-punctate. Claws with c. 6 teeth. Sec-
ondary sexual characters: 6 tarsi as genus;
6 middle tibiae slightly bent in at apex but
not tuberculate-serrate (Fig. 161); 6 with
usually 3, 9 5 or 6 apical ventral setae each
side. Measurements: length 8.5-10.0; width
2.9-3.4 mm.
Types. Holotype $ (British Mus.) and
8 paratypes (3 in M.C.Z., Type No. 31,469)
from Mt. Baduri, Japen Is., West N. G.,
1000 ft. (305 m), Aug. 1938 (Cheesman);
and the following additional paratypes
from West N. G.: 1, R. Manai-Undei,
Japen Is., 500 ft. (c. 150 m), Oct. 1938
(Cheesman); 3, Mt. Lina, Cvclops Mts.,
3500-4500 ft. (c. 1070-1370 mj, Mar. 1936
(Cheesman); 6, Sibil, Star Rge., 1260 m,
dates in May, June 1959 ( Neth. N. G. Exp.,
Leiden Mus.), at hght; 2, Sibil Vy., Star
Mts., 1245 m, Oct. 18-Nov. 8, 1961 (S.
Quate, Bishop Mus.); 1, Bivak 36, Star
Rge., 1220 m, July 29, 1959 (Neth. N. G.
Exp., Leiden Mus.).
Additional material N-E. N. G.: 2,
Eliptamin Vy., 1200-1350 m, June 19-30,
Aug. 1-15, 1959 (W. W. Brandt, Bishop
Mus.); 1, Feramin, 1200-1500 m, June 1-6,
1959 (W. W. Brandt, Bishop Mus.); 1,
Pindiu, Huon Pen., 1200-1450 m, Apr. 18,
1963 (Sedlacek). Papua: 3, Mafulu, 4000
ft. (c. 1220 m), Jan. 1934 (Cheesman); 1,
\\'akaiuna, Sewa Bay, Normanbv Is., Dec.
11-20, 1956 (W. W. Brandt, Bishop Mus.).
Measured specimens. The i holotype and
1 9 paratype from Japen Is.
Notes. D. brunnea resembles forma in
most key characters, but brunnea has oviter
apical elytral angles acutely denticulate (c.
right in forma), 3rd intervals usually 3-
punctate (2-punctate in forma), and S
middle tibiae slightly bent-in but not
tuberculate-serrate as in forma.
Actually, the punctures of the 3rd inter-
vals vary slightly. Two individuals of
brunnea (the holotype and the paratype
horn Bi\ak 36) have 3 punctures on one
and 2 on the other side, although all other
brunnea listed above are 3-punctate on
both sides.
Demetrida fumipes n. sp.
Description. See Plate 2, figure VI; with
characters of genus; form slender, with
prominent eyes and short-spined elytra;
reddish brown, elytra ± paler on disc but
with sides behind humeri blackish, legs
pale with outer edges of tibiae and apices
of femora dark or legs more extensively
dark; not pubescent, reticulate microsculp-
ture faint or light even on elytra, surface
not much punctulate. Head 1.12 and 1.18
width prothorax; eyes prominent, genae
shorter and not prominent. Prothorax
quadrate; width length 1.05 and 0.99; base/
apex 1.21 and 1.17; base/head 0.83 and
170 Biilletin Museum of Companitive Zoology, Vol. 137, No. 1
0.82; sides almost straight or weakly arcuate
in anterior %, sinuate before c. right but
irregular basal angles; margins narrow, each
with seta-bearing puncture before middle,
none at base; surface scarcely punctate
even baso-laterally. Ehjtiri: width elytra
prothorax 2.07 and 2.26; apices short-spined,
outer angles acute, sutural angles slightly
blunted; striae impressed, slightly punc-
tulate; intervals slightly convex, faintly
sparsely punctulate, 3rd usually 3-punctate.
Claws with c. 5 or 6 teeth. Secondary sexual
characters: i tarsi as genus; 6 middle
tibiae slightly bent-in at apex but not
tuberculate-serrate; 6 with 2 or 3, 9 4-6
apical ventral setae each side. Measure-
ments: length 8.3-9.4; width 2.7-3.2 mm.
Tifpes. Holotype S (Bishop Mus.) and
17 paratypes (6 in M.C.Z., Type No. 31,470)
all from Wan and vicinity (including Mt.
Missim), Morobe Dist., N-E. N. G., alti-
tudes from 1100 to 1500-1900 m, dates in
Feb., Mar., Apr., July, Sept., Nov., 1961-
1963 (holotype, Wan, 1200-1300 m, Apr. 6,
1963) (Sedlaceks).
Additional material. Papua: 3, Doveta,
Amazon Bay Dist., 2400 ft. (730 m), Aug.
1962 (W. W. Brandt, C.S.I.R.O.). N-E.
N. G.: 1 teneral, Wau, 1200 m, Sept. 2,
1961 (Sedlacek).
Measured specimens. The S holotype and
1 9 paratype from Wau.
Notes. Among species that are not ob-
viously bicolored, fumipes is closest to
forma but has legs in jiart darker, sides of
elytra behind humeri darker, and 3rd in-
tervals 3-punctate (2-punctate in forma).
Among bicolored species, fumipes is nearest
humcraJis but is smaller, witli humeri less
extensively black. See also comparative
notes under following species.
Demefrida veiafa n. sp.
Description. With characters of genus;
lonn as in preceding species (fumi))es);
reddish brown, disc of elytra ± paler but
sides ol elytra narrowly blackish behind
humcMi, legs in part dark (at least darker
than elytral disc); not pulK\scent, reticulate
microsculpture faintly indicated on pro-
notum and sometimes on part of head and
distinct (but light) on elytra, surface not
much punctulate. Head 1.14 and 1.08
width prothorax; eyes prominent, genae
shorter, not prominent. ProtJiorax quadrate,
long; width length 0.94 and 1.00; base apex
1.21 and 1.34; base head 0.84 and 0.88;
sides weakly irregularly arcuate anteriorly,
strongly sinuate before right or slightly
acute basal angles; margins narrow, each
with seta-bearing puncture before middle
but none at base; baso-lateral areas irreg-
ularly subpunctate. Elytra: width elytra/
prothorax 2.05 and 2.16; apices short-spined
or acutely toothed, with outer angles ± right
and sharply defined, sutural angles obtuse
or slightly rounded; striae impressed, faintly
punctulate; intervals slightly convex, faintly
sparsely punctulate, 3rd usually 2-punctate.
Claws with 5 or 6 teeth. Secondary sexual
characters: i tarsi as genus; i middle
tibiae tuberculate-serrate (c. 6 low tuber-
cles ) ; S with 3 ( rarely 4 ) , 9 c. 5 setae
each side last ventral segment. Measure-
ments: length 8.4-8.8; width 2.7-3.0 mm.
Types. Holotype i ( Bishop Mus. ) from
Saidor, Kiambavi Village, Finisterre Rge.,
N-E. N. G., July 22-29, 1958 (W. \\\
Brandt ) , and 9 paratypes from Finisterre
Rge. (3 in M.C.Z., Type No. 31,471) as
follows: 1, same data as holotype except
Aug. 1-28; 7, Saidor, Ahitoko (N'illage),
Aug. 29-Sept. 5, Sept. 6-24, 1958 (all col-
lected by W. W. Brandt).
Measured specimens. The i holot>pi' and
1 9 paratype from Matoko Village.
Notes. This apparent relati\c> of fornui
is distinguished from the latter b\- dark
elytral edges and daik legs and from
fumipes b\' usualK 2-punctate rather than
3-punctate 3rd intervals, and celata differs
from th(\se and from other similar species
also in Inning ic'ticiilate microsculpture
N'isible, although hiint, on pronotum (and
sometimes part ol head ) as well as elytra.
D. veldid may be more closely related to
diversa but is less distinctK bicolored and
The Carabid Beetles of New Guinea
Darlington
171
more distinctly microreticulate, w ith usually
better developed (but still short) elytral
spines.
The 3rd intervals are 2-punctatc on both
sides of all individuals except that an extra
(3rd) puncture is present on one side only
in two individuals.
One specimen of the type series has
moth scales stuck to it and is presumably
from light-trap material.
Demefrida nigripes n. sp.
Description. With characters of genus;
form as in Figure 106; head and prothorax
red, elytra black, legs and antennae exten-
sively dark with pale bases, tarsi paler; not
pubescent, reticulate microsculpture virtu-
ally absent in i , present (moderateh'
transverse ) on elytra in 9 , surface not
much punctulate. Head 0.98 and 0.94 width
prothorax; eyes prominent, genae shorter
and oblique. Prothorax cordate-subquad-
rate; width length 1.20 and 1.28; base/apex
1.39 and 1.34; base head 0.94 and 0.98;
sides broadly slightly irregularly arcuate in
more than anterior n, moderateh' sinuate
before e. right posterior angles; margins
rather wide, each with seta-bearing punc-
ture slightly before middle but none at
base; disc slightly punctate at sides and
base. Elytra parallel; \\'idth elytra pro-
thorax 1.77 and 1.71; apices with short
spines, outer angles well defined but
slightly obtuse, sutural angles obtuse-
blunted; striae less impressed but more
punctulate than usual; intervals slightly or
not convex, 3rd with 3 dorsal punctures
(middle puncture sometimes doubtful).
Claics with 5 teeth. Secondary sexual char-
acters: i tarsi as genus; 6 middle tibiae
tuberculate-serrate (3 or 4 rounded tuber-
cles); 6 with 2, 9 3 apical ventral setae
each side. Measurements: length 7.4-7.7;
width 2.3-2.7 mm.
Types. Holotype £ (Bishop Mus.) from
Swart Vy., Karubaka, N-E. N. G., 1350 m,
Nov. 18, 1958 (Gressitt); and 1 9 para-
type (Bishop Mus.), Daradae, nr. Javarere,
Musgrove R., Papua, 100 m ?, Oct. 2,
1958 (Gressitt).
Notes. Although the 2 specimens listed
above are from different localities, they
agree in so many ways ( in spite of disagree-
ment in a few details ) that I feel sure they
are conspecific and that they represent an
unusually distinct species, characterized by
form, color, and relatively light but strongly
punctulate elytral striae, as well as by other
characters given in the Key to Species of
Demetrida of New Guinea.
Demefrida dorsalis n. sp.
Description. See Plate 2, figure VII; with
characters of genus; black, elytra with large
common red area centered behind middle,
appendages dark; not pubescent, reticulate
microsculpture absent or indistinct, but
parts of upper surface ( especially elytra )
sparsely punctulate. Head 0.93 and 0.89
width prothorax; eyes prominent, genae
shorter, oblique; front slightly convex, im-
pressed each side anteriorly, slightly punc-
tate at middle. ProtJu^rax subcordate;
width length 1.33 and 1.39; base apex 1.34
and 1.33; base/head 0.95 and 0.96; sides
broadly arcuate in more than anterior %,
strongly sinuate before right or slightly
acute posterior angles; margins rather wide,
each with seta-bearing puncture at or
slightly before middle but none at base;
surface slightly punctate at base and sides.
Elytra: width elytra prothorax 1.77 and
1.78; apices angulate, the angulations right
or slightly acute, outer angles obtuse or
narrowly rounded, sutural angles obtuse;
striae well impressed, scarcely punctulate;
intervals convex, sparsely but distinctly
punctulate, 3rd with c. 4 dorsal punctures.
Chiics with c. 5 teeth. Secondary sexual
characters: S tarsi as genus (squamae of
front tarsi disarranged, but probably in 2
series ) ; S middle tibiae tuberculate-serrate
(c. 4 low rounded tubercles); S with 2 or
3, 9 3 or 4 apical ventral setae each side.
Measurements: length 6.8-8.2; width 2.7-
3.1 mm.
172 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Ti/pes. Holotype S (Bishop Miis.) and
7 paratypes (3 in M.C.Z., Type No. 31,472)
all from Wau and vicinity (inchiding Mt.
Missim and Nami Creek), Morobe Dist.,
N-E. N. G., altitudes from 1100 to 1650
m, dates in Jan., Feb., Mar., 1962, 1963
(holotype. Wan, 1100 m, Jan. 31, 1963)
( Sedlacek ) .
Measured specimens. The i holotype and
1 5 paratype.
Notes. Although known only from a
single locality, this species seems a distinct
on(\ characterized by form, color, angulate
but not spined elytral apices, and virtual
absence of dorsal elytral reticulate micro-
sculpture, as well as by other key char-
acters.
Demetrida basalis n. sp.
Description. With characters of genus;
form and most characters of duplicata ( p.
000); red or brown with base of elytra
(sometimes only humeri) black, legs and
antennae reddish with part of femora and
tibiae usually darker; not pubescent, micro-
sculpture visible (sometimes faint or in-
distinguishable ) only on elytra, surface in
part sparsely punctulate. Head 0.93 and
1.02 width prothorax; eves prominent, genae
shorter, oblique. Prothorax ({uadrate-sub-
cordate; width/length 1.21 and 1.17; base/
apex 1.36 and 1.30; base ^head 0.92 and
0.87; sides irregularly arcuate in more than
anterior 'V\, sometimes subangulate at setae,
strongly sinuate before c. right or slightly
acute sometimes slightly blunted posterior
angles; margins moderately wide, each with
seta-bearing puncture at or slightly before
middle but none at base; disc slightK
punctate across base and in margins.
Ell/Ira: width elytra/prothoiax 1.S6 and
2.10; apices spined, outer angles ± right and
sharply defined, sutmal angles slightly
obtuse, blunted; striae impressed, scarcely
punctulat(\ intervals slightly convex, slightly
punctulate, 3rd with 2 principal and 1 or 2
intermediate smaller dorsal punctures (all
specimens). Clans with r. 5 teeth. Sec-
ondary sexiad chdraclcrs: i tarsi as genus;
4 middle tibiae tuberculate-serrate (c. 6
small tubercles); i with 2 or 3, 9 3 (or
more?) apical ventral setae each side. Mea-
surements: length 8.4-10.0; width 2.9-3.5
mm.
Types. Holotype $ (Bishop Mus.) from
Swart Vv., West N. G., W. ridge, 1800-
2000 m,^ Nov. 19, 1958 (Gressitt); and
paratypes as follows. West N. G.: 4(2 in
M.C.Z., Type No. 31,473), Swart Vy.,
Karubaka, 1500 m, Nov. 11, 20, 1958 (Gres-
sitt); 2, Wissel Lakes, Kamo Vy., Itouda and
Moanemani, 1500-1700 m, Aug. 18, 16, 1962
(Sedlacek); 1, Star Rge., Sibil Vy., 1245
m, Oct. 18-Nov. 8, 1961 (S. Quate, Bishop
Mus.); 1, same locality, 1260 m, June 16.
1959 (Neth. N. G. E.xp., Leiden Mus.), at
hght; 1, Star Rge., Bivak 39, 1300 m, June
28, 1959 (Neth. N. G. Exp., Leiden Mus.);
1 teneral, Araucaria Gamp, 800 m, Apr. 2,
1939 (Toxopeus).
Additional material. One teneral, Karu-
baka, 1450 m, Nov. 16, 1958 (Gressitt),
light trap.
Measured specimens. The 6 holotype and
1 9 paratype from Karubaka.
Notes. This ma>- be a geographic ( west-
ern ) representatix'e of duplicata ( p. 164 )
with elytra black at base rather than en-
tirely reddish brown (but intermediate
color fonns occur as noted under duplicata )
and with elytra Ic^ss distinctb microreticu-
late. Its placc^ among other similarly bi-
colored species is indicated in the Keij to
Species.
Demefrida diverso n. sp.
Description. See Plate 2, figure MIL
with cliaracters ot genus; lorm slender,
with prominent eyi's and acutcK angulate
or short-spined el\ tral apices; color diverse,
brownish red with either whole base of
elytra (except suture ) or onl\ humeri either
black or green (indi\idu;il \ariation), legs
either entireK' red or almost entiicly black
(h()lot\pe, basal '.: ol ehtra black, legs
red); not pubeseint. reticulate microscul]!-
ture indistinc-t or light rwn on elytra, sur-
laee not much piuulnlate. Head 1.07 and
The Carabid Beetles of New Guinea • Darlington 173
1.14 width prothorax; eyes prominent, genae
shorter and obHque. Prothorax subquadrate
or trapezoidal with base varying from nar-
rower than to wider than widtli at middle;
width (at middle) length 1.04 and 0.95;
base apex 1.39 and 1.46; base head 0.88
and 0.94; sides weakly irregularly arcuate
in c. anterior %, sinuate before c. right or
acute but blunted or narrowly rounded
posterior angles; margins rather narrow,
each with seta-bearing puncture before mid-
dle but none at base; surface irregularly
punctate basally and in margins. Elytra:
width elytra/ prothorax (at middle) 1.97
and 2.25; apices with slightly obtuse or
acute angulations or very short spines,
outer angles well defined but varying from
slightly obtuse to acute, sutural angles
obtuse; striae moderately impressed, slightly
punctulate; intervals slightly convex, 3rd
usually 2- sometimes 3-punctate ( see Notes,
below). Claws with c. 5 or 6 teeth. Sec-
ondary sexual characters: 6 tarsi as genus;
S middle tibiae tuberculate-serrate (c. 7
small tubercles); 6 with 2 or 3, 9 4 or
more apical ventral setae each side. Mea-
surements: length 7.5-9.5; width 2.7-3.3
mm.
Types. Holotype c^ (Bishop Mus.) and
27 paratypes (some in M.C.Z., Type No.
31,474) all from Wau and vicinity (in-
cluding Mt. Missim), Morobe Dist., N-E.
N. G., altitudes from 1090 to 1700 m, dates
in Jan., Feb., Mar., May, June, July, Sept.,
Oct., Nov. 1961-1963 (holotype, 1250 m.
May 3, 1963) (Sedlaceks).
Additional material. N-E. N. G.: 3,
Eliptamin Vy., 1200-1350, 1665-2530 m,
June 23-30, July 1-15. 16-31, 1959 (W. W.
Brandt, Bishop Mus.); 1, Korop, Upper
Jimmi Vy., 1300 m, July 12, 1955 (Gres-
sitt), in light trap; 1, Swart Vy., Karubaka,
1550 m, Nov. 8, 1958 (Gressitt, No. 3145);
1, Jim(m)i R., E. Highlands, July-Sept.
1961 (W. W.Brandt, G.S.I.R.O.).
Measured specimens. The S holotype and
1 9 paratype.
Notes. The series from Wau shows the
entire range of variation indicated in the
preceding Description. Occurrence together
of such diverse individuals in what seems
to be one population suggests Mendelian
dimorphism of color (elytral bases black
or green, legs red or black ) , and exceptional
genetic variation of some other characters.
The different characters vary indepen-
dently. For example, leg color is not cor-
related with color or extent of basal elytral
marks. Of the type series, 15 individuals
have 2 punctures on each 3rd interval, 4
(including the holotype) have 2 on one
side and 3 on the other, and 2 individuals
have 3 punctures on each side.
For distinguishing characters of diversa
see the Key to Species of Demetrida of New
Guinea.
Demetrida vigil n. sp.
Description. With characters of genus;
form as in Figure 107; head and prothorax
brownish red, elytra red with basal V.\
black, antennae red, legs mainly black; not
pubescent, reticulate microsculpture faint
and irregular even on elytra in i ( possibU'
more distinct in 9 ), surface not much
punctulate. Head 1.03 width prothorax;
eyes not larger than usual but exceptionally
abruptly prominent, joining neck posteriorly
with virtually no genae. Vrothorax long-
quadrate with anterior angles virtually ob-
literated; width/ length 0.99; base/apex 1.39;
base/head 0.91; sides weakly arcuate,
sinuate before c. right basal angles; margins
narrow, each with seta-bearing puncture
at or slightly before middle (an extra ad-
ventitious puncture on left) but none at
base; disc very convex, slightly rugulose
and punctulate especially in baso-lateral
depressions. Elytra: width elytra prothorax
1.92; apices spined, outer angles acute-
denticulate, sutural angles obtuse-blunted;
striae slightly impressed, faintly punctulate;
intervals slightly convex, 3rd 2-punctate.
Claus with 4 or 5 teeth. Secondary sexual
characters: 6 tarsi as genus; S middle
tibiae weakly tuberculate-serrate (c. 4 low
tubercles); c^ with 3 apical ventral setae
174 Bulletin Muscidu of Comparative Zoology, Vol. 137, No. 1
each side; 9 unknown. Mcasiiiements:
length 7.8; width 2.5 mm.
Type. Holotype S (C.S.I.R.O., Can-
berra) from Doveta, Amazon Bay Dist.,
SE. Papua, 2400 ft. (c. 730 m), Aug. 1962
(W. W. Brandt); the type is unique.
Notes. The eyes, more abruptly promi-
nent than in anv other Denietrida that I
know, distinguish vigil from such similarly
colored species as diversa and divisa.
Demefrido nigriceps n. sp.
Description. With characters of genus;
form ( Fig. 10(S ) e. average, with prominent
but not abrupt eyes, rather narrow pro-
thorax, and spined elytra; head and pro-
thorax black, elytra entirely brown, ap-
pendages brown with antennae darker
outwardly; not pubescent, microsculpture
indicated (faint and irregular) on elytra
only. Head 0.96 and 1.01 width prothorax;
eyes prominent, genae shorter, oblique.
Prothorax subquadrate, long, with base
sometimes wider than middle; width length
0.96 and 1.01; base/apex 1.30 and 1.30; base/
head 0.(S9 and 0.85; sides subparallel in ante-
rior 'Yi, faintly angulate at lateral setae,
broadly sinuate before right or slightly acute
posterior angles; lateral margins narrow,
each with seta-bearing puncture at or
slightly before middle but none at base; sur-
face scarcely or slightly punctate in luargins.
Elytra: width elytra prothorax 2.21 and
2.14; apices spined, outc-r angles acute
and subdenticulate, sutural angles obtuse-
blunted; striae slightly iiupressed, slightly
pimctulate; intervals scarcely convex, 3rd
2-punctate. ('lans with 6 or 7 teeth. Sec-
ondary sexual characters: ', tarsi as genus;
$ middle tibiae not inodificxl ( r. straight,
not tuberculate-serrate, in both s|:>ecimens ) ;
i with 2-4 apical ventral setae each side
(holotype, 4 on each side; paratype, 2 on
one side, 3 on other j; v unknown. Mea-
surements: length e. 10.0; width 3.2 mm.
Types. Holotype S (Bishop Mus.) and
1 ^'paratypc^ (M.C.Z., Type No. 31,475)
both Iroin Sibil Vall(>v, Star Mts.. Wrsi
N. G., 1245 m, Oct. 18-Xov. 8, 1961 (S.
Quate), the holotype at light, the paratype
in Malaise trap.
Notes. The unmodified 6 tibiae distin-
guish this species among other similar ones,
and other differential characters are given
in the Key to Species of Demetrida of New
Giunea.
The sex { i S ) of both specimens has
been determined by dissection as well as
by examination of the front tarsi.
Demetrida saidor n. sp.
Description. With characters of genus;
form slender, with prominent but not
abrupt eyes and spined elytra; head and
prothorax brownish black, elytra brownish
testaceous with humeri and sometimes en-
tire base narrowly black, appendages ir-
regularly dark with paler tarsi; not pubes-
cent, reticulate microsculpture visible ( faint
or light) only on elytra, surface not much
(slightly, finely, sparsely) punctulate. Head
1.25 and 1.11 width prothorax; eyes promi-
nent, genae short, oblique. Prothorax long-
quadrate with relatively wide base; width/
length 0.93 and 1.02; base apex 1.43 and
1.30; base/head 0.85 and 0.88; sides sub-
parallel or slightly arcuate in c. anterior ^^i,
sinuate before usually acute but slightly
blunted posterior angles; margins narrow,
each \\'ith seta-bearing puncture at or
slightly before middle but none at base;
surface slightK' punctate at base and in
margins. Elytra: width ehtra prothorax
2.28 and 2.17; apices spined, outer angles
acute, sharpK defined, sutural angles ob-
tuse; striae lightK impressed, punctulate;
intervals I hit or slightK' comcx, 3rd 2-
punctate (all specimens), the posterior
puncture fai- back. Clatrs with c. 6-7 teeth.
Seconchiry sexii(d characters: S tarsi as
g{>nus; -; middl(> tibiae not or \ery littU^
modilied, not bent in at apex and not
tuberculate-serrate; ,^ with 3, 9 c. 6 apical
ventral setae each side. Measurements:
length 8.6-10,0; width 2.8-3.3 mm.
I'yiics. ?Tolotype S (Bishop Mus.) from
The Carabid Beetles of New Guinea
Darlington
175
Saidor, Matoko Village, Finisterre Rge.,
N-E. N. G., Sept. 6-24, 1958 (W. W.
Brandt); 1 6 paratype (M.C.Z., Type No.
31,476) from Saidor, Kiambavi Village,
Aug. 1-28, 1958 (W. W. Brandt); 2(69)
paratypes from Sepalakembang, Salawaket
Rge., N-E. N. G., 1920 m, Sept. 11-14, 12,
1956 (E. J. Ford, Jr., Bishop Mus.), in
light trap.
Additional material. West N. G.: 1 ten-
eral 6 , Wissel Lakes, Moanemani, Kamo
Vy., 1500 m, Aug. 19, 1962 (Sedlacek).
Measured specimens. The c$ holotype and
the 9 paratype from Sepalakembang.
Notes. Among more or less similar species
(fiimipes, velata, nigriccps) this is distin-
guished by combination of polished ( not
microreticulate) black head and pronotum,
black humeri and legs, and simple 6 middle
tibiae.
Demefrido divisa n. sp.
Description. With characters of genus;
form c. average, with prominent but not
abrupt eyes, rather narrow prothorax,
spined elytra; red or yellowish with basal
% or V-i of elytra black, lower surface red-
dish yellow with metepisterna mainly dark,
legs testaceous with dark knees, antennae
brown; not pubescent, reticulate micro-
sculpture distinct (light) only on elytra,
surface not much punctulate. Head 1.14
and 1.11 width prothorax; eyes prominent,
genae oblique and shorter than eyes, some-
times slightly convex in profile but not
very prominent. Prothorax quadrate, long;
width length 0.95 and 0.95; base apex 1.30
and 1.32; base/head 0.88 and 0.95; sides
nearly straight (except subangulate at
setae) in more than anterior ■'4, sinuate
before c. right or acute, slightly blunted
posterior angles; margins rather narrow,
each with seta-bearing puncture before
middle but none at base; surface vaguely
subpunctate baso-laterally. Elytra: width
elytra prothorax 2.05 and — (elytra spread);
apices variably spined (spines usually short),
outer angles sharph' defined, right or acute,
sutural angles obtuse; striae impressed,
scarcely punctulate; intervals slightly con-
vex, sparsely slightly punctulate, 3rd with
2 or 3 dorsal punctures (see following
Notes). Claws with c. 7 teeth. Secondary
sexual characters: 6 tarsi as genus; S
middle tibiae scarcely modified, at most
slightly bent in at apex, not tuberculate-
serrate; $ with c. 4, 9 5 or 6 apical ventral
setae each side. Measurements: length
9.5-11.5; width 3.1-3.9 mm.
Types. Holotype $ (Bishop Mus.) and
1 <i 'paratype (M.C.Z., Type No. 31,477)
from Tsenga, Upper Jimmi Vy., N-E. N. G.,
1200 m, July 14, 1955 (Gressitt); and addi-
tional paratvpes as follows. N-E. N. G.: 2,
W'au, iMorobe Dist., 1200 m, Sept. 27, 1961,
May 1-15, 1962 (Sedlacek), in light trap;
5, Okapa ( Hornabrook ) ; 1, Swart Vy.,
Karubaka, 1300 m, Nov. 7, 1958 (Gressitt);
1, Sattelberg (British Mus.); Papua: 1,
Kokoda, 1200 ft. (366 m). May 1933
(Cheesman); 2, Dogon, Amazon Bay Dist.,
2400 ft. {c. 730 m), Sept., Oct.-Nov. 1962
(W. W. Brandt, C.S.I.R.O.). West N. G.:
1 9 , Bomberi, Vogelkop, 700-900 m, June
5, 1959 (Gressitt).
Measured specimens. The 6 holotype and
9 paratype from Sattelberg.
Notes. D. divisa resembles one of the
color forms of diver.sa, but divisa is larger,
with elytra at least short-spined and S
middle tibiae not tuberculate-serrate as in
diversa.
In the 6 holotype and S paratype from
Tsenga and also the 9 from Bomberi the
3rd intervals are 2-punctate; in all other
specimens, 3-punctate; but I find no other
characters to suggest that this is a specific
difference.
The specimen from Sattelberg is labeled
by Andrewes, "Genus mihi ignotum."
Demefrida humerolis n. sp.
Description. With characters of genus;
form c. average, with prominent eyes,
quadrate prothorax, and short-spined elytra;
reddish brown, humeri black, legs black or
bicolored; not pubescent, microsculpture
176 Bulletin Muacuni of Coiup<irativc Zoology. Vol. 137, No. 1
faint even on elytra, surtace not much
punctulate. Head 1.07 and l.OS width pro-
thorax; eyes prominent, genae shorter and
obHque. ProtJiorax sul)Ciuadrate with rather
broad base; width length 1.06 and 1.00;
base /apex 1.33 and 1.23; base/head 0.89
and 0.90; sides weakly irregularly arcuate,
usually subangulate at setae, broadly sinu-
ate before right or slightly acute posterior
angles; margins rather narrow, each with
seta-bearing puncture at or before middle
but none at base; disc subpunctate across
base and in margins. Elytra: width elytra
prothorax 1.99 and 2.16; apices short-spined.
outer angles sharply defined, usually dentic-
ulate, sutural angles c. right or slightly
ol)tuse; striae moderately impressed, faintly
punctulate; intervals slightly convex, 3rd
usually 3-punctate. Claws with 5 or 6 teeth.
Secondary sexual characters: 6 tarsi as
genus; i middle tibiae scarcely modified,
at most slightly bent in at apex but not
tuberculate-serrate; 6 with 3, 9 c. 6 apical
ventral setae each side. Measurements:
length 9.3-10,8; width 3.0-3.5 mm.
Types. Holotype 6 (Bishop Mus.) and
12 paratypes (some in M.C.Z., Type No.
31,478) all from Swart Vy., Karubaka, N-E.
N. G., 1300 to 1600 m, dates in Nov. 1958
(holotype, 1300 m, Nov. 7) (Gressitt).
Additional material. N-E. N. G.: 2, Kas-
sem, 48 km E. of Kainantu, 1350 m, Nov. 7,
1959 (T. C. Maa, Bishop Mus.); 1, Tsenga,
Upper Jimmi Vy., 1200 m, July 13, 1955
(Gressitt); 1, Jim(m)i R., E. Highlands,
Julv-Sept. 1961 (W. W. Brandt, C.S.I.R.O.).
West N. G.: 1, Sibil, Star Rge., 1260 in.
May 24, 1959 (Neth. N. G. Exp., Leiden
Mus. ) .
Mea.sured s])eciniens. The •; holot\pc and
1 9 paratyjie from Karubaka.
Notes. P. huntcralis differs Irom the
preceding species (divisa) i)riucipall\ in
color, having less black on eUlral bas(>s
but darker legs. It is close also to fut)ti))es
but is larger and more lu>a\ily marked.
The interr(4ationships ol these forms are
still not clear.
The 3rd inter\als are usually .3-punetate
in humeralis but are only 2-punctate in the
indixidual from Sibil.
Demefrida imitafrix n. sp.
Description. See Plate 3, figure IX; with
characters of genus; relatively wide; dark
blue-black with dark appendages; not
pubescent, reticulate microsculpture absent
or faint, but surface finely sparsely punc-
tulate. Head 0.84 and 0.88 width prothorax;
eyes prominent, genae much shorter and
oblicpie. Frothorax subcordate; width
length 1.39 and 1.36; base apex 1.36 and
1.40; base head 1.02 and 1.02; sides
broadly slightly irregularly rounded an-
teriorly, strongly sinuate before c. right
slightly l)lunted posterior angles; margins
moderately wide, each with seta-bearing
puncture at or slightly before middle but
none at base; disc subpunctate baso-later-
ally. Elytra short and wide; width ebtra
prothorax 1.74 and 1.80; apices spined, outer
angles obtuse or blunted, sutural angles
obtusely blunted; striae impressed, slighth'
or scarcely punctulate; intervals nearly flat
or slightly convex, 3rd 2-punctate. Claics
with 4 or 5 teeth. Secondary sexual cliar-
acters: 6 tarsi as genus; o middle tibiae
with inner edge swollen or thickened before
apex, the swollen portion separated from
the apex b\' a broad emargination; i with
2 or 3, 9 c. 4 apical ventral setae each side.
Measurements: length 7.4-8.5; width 3.0-
3.5 mm.
Types. IIolotN'iie i (Bishop Mus.) from
Karimui, S. of Goroka, N-E. N. (i., 1000 m,
Inne 2, 1961 (Gressitt), taken in light trap;
and paratopes as follows. Papua: 1 broken
V, Dobodura, Mar.-Jul\ 1944 (Darlington)
(M.G.Z., Type No." 31,479), taken on a
lighted window; 1, Kokoda, 1200 ft. (366
m), Aug. 1933 (Gheesman); 1. Dogon,
Amazon Bay Dist., 2400 ft. ( r. 730 m).
Sept. 1962 (W. W. Brandt, C;.S.1.R.(). ); 1,
Misima Is., No\ . 1963 (\V. \V. Brandt,
C:.S.I.R.O.). West N. <;.: 1. Gamp 2,
Sabron. Gyclops Mts., 2000 ft. (610 m),
|nl\ 1936 ((>heesman).
The Carabid Beetles of New Guinea
Darlington
177
Measured specimens. The 6 holotype and
the 9 paratype from Dogon.
Notes. Among the New Guinean species
of Demetricla, this one is unique in its
broad form and in form of i tibiae. Never-
theless it has the essential characters of
Demetrida and I do not think it should be
separated from that genus, at least not
unless the genus as a whole is divided.
Superficially, D. imitatrix resembles Vio-
higonum violaceiim (Chaudoir), which
is very common at low altitudes in New
Guinea. This may be an example of
Batesian mimicry.
Demefrida viridipennis n. sp.
Description. See Plate 3, figure X; with
characters of genus; not pubescent; head
and prothorax red, elytra bright green usu-
ally shading to purple toward apex, ap-
pendages reddish yellow; reticulate micro-
sculpture visible ( light ) only on elytra, but
surface in part slightly sparsely punctu-
late. Head 1.08 and 1.11 width prothorax;
eyes moderately prominent, genae shorter
and oblique, not prominent. Prothorax
subquadrate; width length 1.08 and 1.03;
base/ apex 1.32 and 1.31; base head 0.88
and 0.86; basal angles c. right, c. blunted;
margins rather narro\\', each \\\i\\ seta near
or slightly liefore middle but none at base;
disc subpunctate across base and in margins.
Elytra: width elytra/prothorax 1.99 and
2.17 (latter spread by pin?); apices spined,
outer angles sharply defined, c. right ( some-
what variable), sutural angles obtuse; striae
deeply impressed, slightly punctulate; in-
tervals slightly convex, faintly punctulate,
3rd with 2 dorsal punctures. Claws with c.
5 teeth. Secondary sexual characters: S
tarsi as genus; c^ middle tibiae tuberculate-
serrate (c. 6 low tubercles); c5 with 2, $
3 setae each side last ventral segment.
Measurements: length 6.9-8.5; width 2.5-
3.1 mm.
Types. Holotype $ (Bishop Mus.) from
Wau, Morobe Dist., N-E. N. G., 1200-1300
m. Mar. 14, 1963 (Sedlacek); and para-
types (some in M.C.Z., Type No. 31,480)
as follows. N-E. N. G.: 4, Wau, 1200,
1220-1250, 1300 m, Nov. 12, 1961, Feb. 11,
Jan. 23, 1963, Oct. 14, 1965 (Sedlaceks);
1, Bulolo (near Wau), 1005 m, Aug. 25,
1956 (E. J. Ford, Jr., Bishop Mus.); 1,
Mt. Missim, 1600-2000 m, Sept. 21-24, 1964
(M. Sedlacek); 1, Karimui, 1080 m, Julv
14-15, 1963 (Sedlacek); 1, Okapa, Apr. 19,
1965 ( Hornabrook ) ; 1, Kainantu, July 9
(Sedlaceks); 1, \\\un. Upper Jimmi Valley,
840 m, July 17, 1955 (Gressitt); 1, Finis-
terre Rge., Saidor, Funvende, 1200 m, Sept.
24-30, 1958 (W. W. Brandt, Bishop Mus.);
1, Adalbert Mts., Wanuma, 800-1000 m,
Oct. 26, 1958 (Gressitt). Papua: 1, Ko-
koda, .200 ft. (366 m), June 1933
(Cheesman); 2, Owen Stanley Rge., Goilala
(Loloipa, Nov. 25-Dec. 10^ and Tapini,
975 m, Nov. 16-25, 1957) (W. W. Brandt,
Bishop Mus.); 1, Mt. Lamington, 1300-
1500 ft. (c. 400-460 m) (C. T. McNamara,
S. Australian Mus.).
Measured specimens. The i holotype and
1 9 paratype from Wau.
Notes. Form, color, and deep elytral
striae set this distinct species off from all
others of the genus kno\\'n to me. It is
evidently widely distributed in at least the
eastern half of New Guinea at moderate
altitudes.
Demetrido lepida n. sp.
Description. See Plate 3, figure XI; with
characters of genus; head and prothorax
black, elytra green-purple (variable, often
more green to\\'ard base and more purple
toward apex, sometimes slightly reddish on
disc), appendages dark; not pubescent,
reticulate microsculpture faint even on
elytra, surface not much punctulate. Head
1.24 and 1.12 width prothorax; eyes rather
abruptly prominent, genae c. long as eyes,
slightK' convex in outline but not very
prominent; front wide, irregularly flattened
and impressed or subpunctate at middle.
Prothorax subquadrate; width length 0.95
and 1.05; base apex 1.30 and 1.22; base/
head 0.74 and 0.77; sides arcuate anteriorly,
sinuate before c. right but blunted or nar-
178 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
rowly rounded posterior angles; margins
narrow, each with seta-bearing puncture at
or shghtly before middle but none at base;
disc convex, baso-lateral impressions almost
obsolete, surface faintly subpunctate across
base and in margins. Elytra: width elytra
prothorax — (elytra spread) and 1.95; apices
long-spined, outer angles acute or denticu-
late, sutural angles obtuse; striae lightly
impressed, lightly punctulate; intervals flat
or slightly convex, .3rd 3-punctate. Claws
with c. 7 teeth. Secondary sexual char-
acters: S tarsi as genus; S middle tibiae
slightly tuberculate-serrate {c. 4 spaced
tubercles); S with 3, 9 4-6 apical ventral
setae each side. Measurements: length
9.2-10.8; width 3.0-3.3 mm.
Types. Holotype 9 ( Bi.shop Mus. ) and
10 paratypes (some in M.C.Z., Type No.
31,-1(S1) from Swart Vy., Karubaka, N-E.
N. G., altitudes from 1300 to 1600 m, dates
in Nov. 1958 (holotype, 1450 m, Nov. 12)
(Gressitt); and additional paratypes as
follows. West N. G.: 17, Wissel Lakes,
Enarotadi, altitudes from 1750 to 1900 m,
dates in July, Aug. 1962 (Sedlacek); 1,
W'isscl Lakes, Itouda, Kamo Vv., 1500-1700
m, Aug. 18, 1962 (Sedlacek); 1, Wissel
Lakes, Kamo-Debei div., 1700 m, Aug. 13,
1955 (Gressitt); 1, Lower Mist Camp,
1700 m, Jan. 17, 1939 (Toxopeus). Papua:
1, Owen StanlcN' Hge., Goilala, Loloipa,
Feb. 1-15, 1958' (W. W. Brandt, Bishop
Mus.).
Measured specimens. A 6 paratype from
Swart Valley and the 9 holotype.
Notes. The bright color, rather abruptly
prominent eyes, and long elytral spines
characterize this fine species. It appears to
be widely distributed in New Guinea at
moderate altitudes. Ot the 19 specimens
seen, only 3 are i $ .
Two Karubaka individuals and one from
JMiarotadi are labe](>(l as taken in light
traps.
Demefrida sublepida n. sp.
Description. With characters of genus;
form r. as in preceding species {lepida)
but eyes less abruptly prominent and elytral
spines shorter; head and prothorax green,
elytra green-purple (variable); appendages
dark, tarsi paler; not pubescent, reticulate
microsculpture visible (faint) only on elytra,
surface not much punctulate. Head 1.16
and 1.11 width prothorax; eyes prominent
but not abrupt, genae shorter and oblique.
Prothorax subquadrate; width length 1.01
and 1.03; base apex 1.20 and 1.18; base/
head 0.80 and 0.86; sides weakly slightly
irregularly arcuate in more than anterior
■'4, usuallv stronglv sinuate before usuallv
acute posterior angles; margins narrow, each
with seta-bearing puncture slightly before
middle but none at base; baso-lateral im-
pressions subobsolete, disc slightly trans-
versely wrinkled, vaguely subpunctate
across base and in margins. Elytra: width
elytra prothorax 2.11 and 2.19; apices
spined, outer angles c. right or obtuse,
sutural angles obtuse; striae well impressed,
scarcely punctulate; intervals convex, finely
sparsely punctulate, 3rd 2-punctate. Cdaws
with 5 or 6 teeth. Secondary sexual cJiar-
acters: i unknown; 9 with 3 or 4 apical
ventral setae each side last ventral segment.
Measurenients: length 7.0-9.0; width 2.5-
3.1 mm.
Tyj)es. Holotype 9 (Bishop Mus.) from
Wissel Lakes, Enarotadi, West N. G., 1850
m, Aug. 1, 1962 (Sedlacek); and addi-
tional paratypes as follows. West N. (i.: 7
(some in \LC.Z., Type Xo. 31,482), Enaro-
tadi, 1750 to 1900 m, dates in July, Aug.
1962 (Sedlacek); 1, Wissel Lakes, IVapura,
Kamo Vv., 1530 m, Aug. 11, 1955 (Gressitt).
N-E. N.'g.: 1, Swart Vy., Karul)aka, 1300
m, Nov. 7, 1958 (Gressitt); 1, Wan, Morobe
Hist., 1.300 m. June 15, 1961 (Gressitt),
on Pii:)turus. All specimens are 9 9 .
Meastircd six'cinwns. Tlu' 9 holotxpe and
9 paratype from Enarotadi.
Soles. Although perhaps related to the
preceding spc^cies (lepida), sid)lcpid(i differs
in a surjirising number ol characters in-
cluding less abrupt eyes, head and pro-
notum green rather tlian black. el\ tral
spines slioitei-, elytial striae deeper, claws
The Carabid Beetles of New Guinea • Darlington 179
with fewer teeth, and size smaller. These
species evidently occur together at some
localities, and they may be involved in some
sort of mimicry.
Demetrida viridibasis n. sp.
Description. With characters of genus;
form slender, with ± prominent eyes and
short-spined elytra; red, c. basal Va of elytra
bright green with the green color extending
back more at sides than at middle, femora
and parts of tibiae dark; not pubescent,
reticulate microsculpture usually visible
(but light) on front of head and on pro-
notum as well as on elytra, much of upper
surface also sparsely finely punctulate.
Head 1.05 and 1.02 width prothorax; eyes
prominent, genae c. long as eyes and
slightly convex in outline but not very
prominent. Frothorax subquadrate, long;
width/length 1.00 and 1.09; base/apex 1.25
and 1.23; base/head 0.85 and 0.85; sides
nearly straight for much of length except
slightly subangulate at setae, broadly sinu-
ate before c. right slightly blunted posterior
angles; margins narrow, each with seta-
bearing puncture slightly before middle but
none at base; disc less convex and with
more distinct baso-lateral impressions than
in k'pida and suhJepida, subpunctate across
base and in margins. Elytra moderately
long; width elytra prothorax 1.98 and 1.86;
apices short-spined, outer angles c. right,
sharply formed, sutural angles obtuse; striae
impressed, faintly punctulate; intervals con-
vex, 3rd 3-punctate. Cdaics with c. 5 teeth.
Secondary sexual characters: i see Notes,
below; 9 with 5 or more apical ventral
setae each side. Measurements: length
8.5-10.3; width 2.9-3.5 mm.
Types. Holotype 9 (C.S.I.R.O., Can-
berra ) from Dogon, Amazon Bay Dist., SE.
Papua, 2400 ft. (c. 730 m), ^Sept. 1962
(W . \V. Brandt); 1 9 paratype (C.S.I.R.O.)
with same data except collected Oct.-Nov.;
1 9 paratype (M.C.Z., Type No. 31,483),
Doveta, Amazon Bay Dist., 2400 ft. (c. 730
m), Aug. 1962 (W. W. Brandt).
Additional material. N-E. N. G.: 3,
Swart Vy., Karubaka, 1300, 1500 m, Nov. 7,
11, 20, 1958 (Gressitt); 1, Finisterre Rge.,
Saidor, Kiambavi Village, Aug. 1-28, 1958
(W. W. Brandt, Bishop Mus.). West N. G.:
1, Cyclops Mts., 3400-4500 ft. (c. 1040-
1370 m). Mar. 1936 (Cheesman).
Measured specimens. The 9 holotype and
9 paratype from Doveta.
Notes. This species seems close to the
green-marked form of diversa but has
longer elytral spines. Some specimens
listed under Additional material are doubt-
fully identified. Most are 9 9 ; the only S ,
from Swart \'y., has middle tibiae slightly
bent in toward apex but not tuberculate-
serrate, and c. 4 apical ventral setae each
side.
Demetrida sibil n. sp.
Description. With characters of genus;
form slender, with prominent eyes and
spined elytra; head, prothorax, and c. basal
Vi of elytra dark greenish, the dark color
extending farther back at sides of elytra
than at middle, and suture sometimes red
almost to base, rest of elytra red, femora
and outer edges of tibiae greenish black,
antennae brown, lo^^'er surface greenish
black in anterior half, abdomen red; not
pubescent, reticulate microsculpture faint
even on elvtra. Head 1.19 and 1.10 width
prothorax; eyes moderately abruptly promi-
nent, genae nearly as long as eyes, oblique;
front flattened, irregularly slightly im-
pressed and subpunctate at middle. Pro-
thorax subquadrate; width length 0.94 and
0.98; base apex 1.38 and 1.28; base head
0.81 and 0.82; sides weakly irregularly
arcuate for much of length, rather abruptly
sinuate before right or slightly acute some-
times slightly blunted posterior angles;
margins narrow, each with seta-bearing
puncture slightly before middle but none at
base; disc strongly convex, with baso-lat-
eral impressions weakly indicated, surface
slightly punctate across base and in margins.
Ehjtra: width elytra/ prothorax 2.09 and
1.98; apices spined, outer angles acute or
denticulate, sutural angles obtuse; striae
180 Bulletin Museum of Co7y}parative Zoology, Vol. 137, No. 1
moderately impressed, finely punetulate; in-
tervals slightly convex, very sparsely in-
conspicuously punetulate, 3rd 3-punctate
(except intermediate puncture lacking on 1
side in 1 paratype). CAciws with c. 6 teeth.
Secondary sexual character: 6 tarsi as
genus; 6 middle tibiae bent-in at apex but
not tuberculate-serrate; c5 with apparently
4, 9 c. 8 or 9 apical ventral setae each side.
Measurements: length 9.4-10.8; width 3.0-
3.4 mm.
Types. Holotype $ (Leiden Mus.) and 5
paratypes (2 hi M.C.Z., Type No. 31,484)
from Sibil, Star Rge., West N. G., 1260 m,
dates in May and June, 1959 (holotype,
June 17) (Neth. New Guinea Exp.), taken
at light; and additional paratypes as fol-
lows. West N. G. : 2, preceding locality
("Star Mts. Sibil Val."), 1245 m, Oct. 18-
Nov. 8, 1961 (S. Quate, Bishop Mus.), at
light. N-E. N. G.: 1, Feramin, 1200-1500
m, June 15-18, 1959 (W. W. Brandt,
Bishop Mus.).
Measured specimens. The i holotype and
1 9 paratype from Sibil.
Notes. The color (head and pronotum as
well as elytra! bases green ) and long-spined
elytra distinguish this from other species of
the diversa complex.
Demetrida seticollis n. sp.
Description. With characters of genus;
form as in Figure 109; brown, with head
and prothorax and sometimes base of elytra
slightly darker; appendages brown; not
pubescent, reticulate inicrosculpture visible
(but very light) only on elytra. Head 1.20
and 1.19 width prothorax; eyes \('r\' promi-
nent, genae {)bli(|ue and not sharply distinct
from neck; front irregularly flattened or
impressed and subpunctate at middle. Pro-
thorax (piadrate-trapezoidal; width /length
1.02 and 0.97; base apex 1.45 ;uid 1.29;
base/head 0.86 and 0.90; sides wcakK
arcuate or c. straight in "'i or more ol length,
strongly sinuate beh)re prominent, c. light
or acute (somewhat varia])le ) basal angles;
margins narrow, each \\ itli seta before mid-
dle and at base and additi<)n;il iisualK
smaller setae anteriorly; disc moderately
convex, with irregular baso-lateral impres-
sions, surface irregular or subpunctate
across base and in margins. Elytra ample;
width elytra/prothorax 2.20 and 2.22; apices
long-spined, outer angles acute-denticulate,
sutural angles obtuse; striae lightly im-
pressed, finely punetulate; intervals slightly
convex or c. flat, 3rd 3- or 4-punctate ( vari-
able, sometimes unsymmetric). Claws with
6-8 teeth. Secondary sexual characters: $
tarsi as genus; i middle tibiae tuberculate-
serrate ( c. 4 widely sometimes irregularly
spaced tubercles); 6 with 2-A, 9 c. 6
apical ventral setae each side. Measure-
ments: length 8.7-10.5; width 2.9-3.4 mm
( except 1 i from Wissel Lakes, doubtfully
identified, 11.3 X 3.7 mm).
Types. Holotvpe 6 ( Bishop Mus. ) from
Wissel Lakes, Enarotadi, West N. G., 1900
m, Aug. 21, 1955 (Gressitt); 61 paratypes
(some in M.C.Z., Type No. 31,485) from
the Wissel Lakes area (Enarotadi, Moane-
mane, Itouda, Urapura, Okaitadi, "Paniai-
Kamo div."), 1500-2050 m, dates in Jul\-,
Aug., 1955, 1962 (Gressitt, Sedlacek );" and
4 additional paratvpcs from the same area,
Arabu Camp, 1800 m, Oct. 7, 8, 12, 17,
1939 (H. Boschma, Leiden Mus.).
Additional material. West N. G. : 1 very
large 6 , data as holot)pe except 1500 m,
Aug. 14, 1962 (Sedlacek); 2, Juliana Bivak,
1800 m, Aug. 30, Sept. 5, 1959 (Neth. N. G.
Exp., Leiden Mus.); 1, Star Rge., Bivak
39A, 1500 m, July 2, 1959 (Neth. N. G.
Exp., Leiden Mus.); 1, Swart Vv., \^^ ridge,
1800-2000 m, Nov. 19, 1958' (Gressitt).
N-i:. ^, (;.: l, Gewak, Salawaket Hge.,
1530 ni, Sept. 6, 1956 ( E. J. Ford. Jr..
Bishop Mus.), in light trap. Papua: 1,
Owen Stanle\- Hge., Cioilala, Bom(\ 1950 m.
Mar. 8-15, 1958 ( W. \\ . Brandt, Bishop
Mus.).
Measured specimens. The S holotype and
I 9 paratype from liinarotadi.
Notes. The extra seta of the prothoracic
margins antciioily distinguish this species
hoin all other noiniubescent Demetrida
known lo ww. These setae aic much
The Carabid Beetles of New Guinea • Darlington 181
stronger and more erect than the fine extra
marginal hairs of D. seriata and niibicola.
When the setae are ]:)roken off, or perhaps
lacking in aberrant individuals, the species
is still recognizable by form especially of
prothorax, color, and long elytral spines.
D. scticoUis apparently ranges widely in
New Guinea at considerable altitudes (not
known below 1500 m ) on the higher moun-
tain ranges. The fact that it has not been
found on the Morobe Plateau (Wau, etc.)
is noteworthy.
Demefrida pollipes n. sp.
Description. With characters of genus;
form slender, with moderately prominent
eyes and strongly spined elytral apices; head
and pronotum reddish piceous, elytra
blackish with small discal area usually
reddish, legs testaceous, antennae brown,
lower surface dark with metepisterna paler;
not pubescent, reticulate microsculpture
absent or indistinct, surface not much punc-
tulate. Head 1.14 and 1.16 width prothorax;
eyes prominent, genae shorter, oblique.
P rot J} o rax subquadratc, long; width length
1.03 and 0.96; base apex 1.35 and 1.34;
base/head 0.84 and 0.86; sides very weakly
arcuate in c. anterior %, strongly sinuate
before right or acute usually slightly
blunted posterior angles; margins narrow,
each with seta-bearing puncture before
middle and at basal angle; disc with mod-
erate baso-lateral impressions, scarcely
punctate. Elytra: width elytra /prothorax
2.04 and 2.05; apices strongly spined, outer
angles acute or denticulate, sutural angles
obtuse; striae lightly impressed, finely ir-
regularly punctulate; intervals almost flat,
3rd usually 2-punctate, sometimes 3-punc-
tate on 1 side. Claws with c. 5 teeth. Sec-
ondary sexual characters: £ tarsi as genus;
S middle tibiae weakly tuberculate-serrate
(c. 4 low tubercles); S with 3, 9 5 or 6
apical ventral setae each side. Measure-
ments: length 8.4-9.6; width 2.5-2.9 mm.
Types. Holotype S (Bishop Mus.) and
19 paratypes (some in M.C.Z., Type No.
31,486) from Wau, Morobe Dist., N-E.
N. G., altitudes from 1100-1500 m, dates
in Jan., Feb., Mar., Apr., Sept., Dec, 1961-
1966 (holotype, 1450 m, Feb. 6, 1963)
(Sedlacek, 1 paratype T. C. Maa); 1 para-
type, Mt. Missim,' 1600-2000 m, Sept. 21-
24, 1964 (M. Sedlacek).
Measured specimens. The S holotype and
1 9 paratype.
Notes. See Key to Species for place of
D. pallipes among other New Guinean
Dcmetrida.
Demetrido discoidalis n. sp.
Description. With characters of genus;
form c. as in preceding species i))allipes);
black, elytra with large elongate common
red area centered behind middle, lower
surface and appendages dark; not pubes-
cent, reticulate microsculpture absent or
indistinct on elytra, surface not much
(sparsely inconspicuously) punctulate. Head
1.06 and 1.13 width prothorax; eyes promi-
nent, genae shorter, oblique. Frothorax sub-
quadrate, but anterior angles rounded to
neck; width length 1.04 and 1.00; base/
apex 1.51 and 1.47; base/head 0.90 and
0.91; sides arcuate through much of length,
sinuate before c. right slightly blunted pos-
terior angles; margins narrow, each with
seta-bearing puncture before middle and at
base; disc with baso-lateral impressions
weak, surface slightly irregular or sub-
punctate across base and in margins.
Elytra: width elytra/prothorax 1.91 and
2.02; apices ( variably ) spined, outer angles
sharply defined, varying from acute to
slightly obtuse, sutural angles obtuse;
striae lightly impressed, punctulate; inter-
vals flat, 3rd 2-punctate. Claws with 6 or
7 teeth. Secondary sexual characters: S
tarsi as genus; 6 middle tibiae tuberculate-
serrate (c. 4 low rounded tubercles); S
with 2 or 3, 9 4 or 5 apical ventral setae
each side. Measurements: length 9.0-11.5;
width 3.0-3.7 mm.
Types. Holotype S (Bishop Mus.) from
Sibii Vy., Star Rge., West N. G., 1245 m,
182 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Oct. 18-Nov. 8, 1961 (S. & L. Quate), in
Malaise trap; and paratypes as follows.
West N. G.: 1, Sibil, Star Rge., 1260 m,
June 1959; 2, Bivak 36, Star Rge., 1220 m,
July 28, 1959; 1, Bivak 39A, Star Rge.,
1550 m, July 5, 1959 (these 4 paratypes all
Neth. N. G. Exp., Leiden Mus.)'. N-E.
N. G.: 1, Eliptamin Vv., 1665-2530 m,
June 19, 1959 (W. W. Brandt, Bishop Mus.);
1, Feramin, 1200-1500 m, June 15-18, 1959
(W. W. Brandt, Bishop Mus.). (Some
paratypes in M.C.Z., Type No. 31,487. )
Measured specimens. The S holotype and
9 paratype from Bivak 39A.
Notes. This may be a geographic repre-
sentative of the preceding species ( paUipes)
from which it differs only slightly in form
but more in color, with larger red area on
elytra and dark rather than pale legs. It
resembles dorsaJis in color but differs in
form (much narrower than dorsaJis), pres-
ence of posterior-lateral prothoracic setae,
and in other ways: in fact, these 2 species
are not closely related.
Demetrida sedlacekorum n. sp.
Description. With characters of genus;
form slender, c. as in preceding species
{paUipes, discoidaJis) ])ut elytra short-
spined; black with bluish tone especially
on elytra, appendages dark; not pubescent;
reticulate microsculpturc> absent or indis-
tinct, but upper surface in part with \er)'
fine, sparse, inconspicuous punctulation.
Head 1.14 and 1.08 width prothorax; eyes
prominent, genae shorter, oblique. Pro-
thorax subcjuadrate, long, with rathcM- broad
base; width length 0.98 and 0.99; base apex
1.41 and 1.43; base/head 0.86 and 0.87;
sides weakly arcuate in c. Ti of length,
sinuate before c. right but variabl(\ blunted
posterior angles; margins rather narrow,
each with seta-bearing imncturc^ before
middle and at l:)asal angle; disc with baso-
lateral impressions deep but small, sub-
|)unctat('. i'Jijtrd: width el\tra prothorax
2.03 and 1.93; apices short-spined, outer
angles sharply defined, usually acute, su-
tural angles obtuse; striae lightly impressed.
punctulate; intervals flat or slighth- convex,
third usually 2-, sometimes 3-punctate.
Claws with 6 or 7 teeth. Secondary sexual
characters: S tarsi as genus; i middle
tibiae weakly tuberculate-serrate (margin
wavy); i with 2 or 3, 9 4 or 5 apical
ventral setae each side. Measurements:
length 8.5-9.8; width 2.6-3.1 mm.
Types. Holotype c^ (Bishop Mus.) and
28 paratypes (some in M.C.Z., Type No.
31,488) all from Wau, Morobe Dist., N-E.
N. G., altitudes from 1180 to 1500 m, dates
in Jan., Feb., Mar., Apr., June, Sept., Nov.,
1961-1964 (holotype, 1220-1250 m, Jan. 23,
1963) (Sedlaceks).
Additional material. N-E. N. G.: 2,
Jim(m)i R., E. Highlands, July-Sept., 1961
"(W. W. Brandt, C.S.I.R.O.). Papua: 1,
Ow^en Stanlev Rge., Goilala, Tororo, 1560
m, Feb. 21-24, 1958 {W . W. Brandt,
Bishop Mus.).
Measured specimens. The i holotype and
1 9 paratype.
Notes. See final couplets of Key to
Species for place of sedlacekortim among
other New Guinean Demetrida.
Demetrida brandti n. sp.
Description. See Plate 3, figure XII;
with characters of genus; form c. of paUipes
and discoidalis, slender, with long-spined
elytra; color entirely bhu^-black. with dark
appendages; not pubescent, microsculpture
\irtuallv absent even on elvtra, surface not
much (very finely, sparsely, inconspicu-
ously) punctulate. Uead 1.09 and 1.07
width prothorax; c\es prominent, genae
shorter, oblicine. Prothorax ({uadrate, long;
width ItMigth 0.98 and 1.03; base ;ipex 1.44
and 1.43; base head 0.87 and 0.88; sides
weakly arcuate in c. ant(Mior 'H, broadly
sinuate before c. right or slightK" acute
posterior angles; margins rather narrow
each with seta near or before middle and
at base; baso-l;iteral impressions moderate,
subpunctate. Elytra: width elytra pro-
thorax 1.98 and 1.94; apices with moderately
long spines, outer angles wc-ll defined, ±
ritiht, sutural angles blunted; striae sli<j:htl\
The Carabid Beetles of New Guinea • Daiiin<'toii
183
impressed, faintly punctulate; intervals
slightly convex, 3rd 2-punctate. Claws with
c. 6 teeth. Secondary sextial characters: i
tarsi as genus; 6 middle tibiae tuberculate-
serrate ( c. 4 tubercles ) ; i with 3, 9 4 or 5
apical ventral setae each side. Measure-
ments: length 8.8-10.2; width 2.8-3.3 mm.
Types. Holotype i (Bishop Mus.) and
2 paratypes from Finisterre Rge., Saidor,
Kiambavi Village, N-E. N. G., 1400 m,
July 22-29 (holotype), Aug. 1-28 (para-
types); 1 paratype, Saidor, Funyende, 1200
m, Sept. 24; 2 paratypes, Saidor, Matoko,
Aug. 29-Sept. 5, Sept. 6-24 (all collected
1958 by W. W. Brandt for Bishop Mus.;
some paratypes now in M.C.Z., Type No.
31,489).
Additional material. N-E. N. G.: 2, Swart
Vy., Karubaka, 1500 m, Sept. 20, 1958
(Gressitt); 1, Gewak, Salawaket Rge., 1530
m, Sept. 6, 1956 (E. J. Ford, Jr., Bishop
Mus.), in light trap. West N. G.: 2,
Wamena, 1700 m, Feb. 10-25, 1960 (T. C.
Maa, Bishop Mus.). Papua: 1, Purosa
Camp, Okapa area, 1950 m, Sept. 23,
1959 (L. J. Brass, Sixth Archbold Exp.,
A.M.N.H.).
Measured specimens. The i holotype and
1 9 paratype from Kiambavi.
Notes. This will probably prove to be a
geographic subspecies of sedlacekorum dis-
tinguished mainly by longer elytral spines.
Genus PHLOEOCARABUS Macleay
Macleay 1871, Trans. Ent. Soc. New South Wales
2, p. 85.
Sloane 1898, Proc. Linnean Soc. New South Wales
23, p. 499.
Csiki 1932, Coleop. Cat., Caraliidae, Harpalinae 7,
p. 1488 (see for additional references, synonymy,
and list of species ) .
Diagnosis. See Key to Genera of Lebiini
of New Guinea and Figure 110.
Description ( characters common to the
2 New Guinean species). Form c. as in
Figure 110; color diverse; not pubescent.
Head: eyes large, prominent; 2 setae over
each eye; antennae pubescent from middle
of 4th segments; front with long, slightly
curved costa on each side passing inside
position of anterior seta; clypeus transverse,
1-setose each side; labrum wide, arcuate-
truncate, 6-setose; mentum with long, entire
tooth; ligula subtruncate with 2 principal
setae, paraglossae attached to ligula, nar-
rowed and rounded to apex of ligula; palpi
rather short, apical segments of labial palpi
widened, c. triangular. Prothorax transverse,
arcuately narrowed anteriorly, slightly lobed
at base; margins rather wide, flat, scarcely
reflexed, each with usual 2 setae; disc with
impressed middle line and weak transverse
impressions; base with fine marginal line
entire or nearly so, apex not margined at
middle. Elytra with rounded, slightly nar-
rowed humeri; apices obliquely sinuate-
truncate, with outer angles broadly and
inner angles narrowly rounded; striae entire,
moderately impressed, not punctate; in-
tervals not specially elevated at base, 3rd
2-punctate with punctures before middle
on outer edge and behind apical % near
inner edge. Inner icings full. Legs mod-
erate; 4th segments middle and hind tarsi
emarginate; 5th segments with accessory
setae; claws each with c. 4 rather long
teeth. Secondary .sexual characters: i front
tarsi scarcely dilated, 2-seriately squamu-
lose; i middle tarsi also squamulose; i
middle tibiae not excised; 6 with 1 prin-
cipal (sometimes a 2nd smaller), 9 2 setae
each side last ventral segment.
Type species. P. mastersi Macleay, of
Australia.
Generic distribution. Australia, with 1
Australian species extending to New
Guinea and New Britain, and an addi-
tional species endemic in New Guinea.
Notes. The 2 species here assigned to
Phloeocarahus are very different super-
ficially but share the technical characters
of the genus.
Key to Species of Phloeocarabvs of
NE^v Guinea
1. Color black or piceous, elytra with testa-
ceous marks (p. 184) nigricollis
- Strikingly bicolored, head and prothorax red,
elytra bine (p. 184) -— euplenes
184 Bulletin Museuni of Comparative Zoology, Vol. 137, No. 1
Phloeocarobus nigricoHis (Macleay)
Macleay 1864, Trans. Ent. Soc. New South W'ak-s
1, p. Ill {Trigoiwthoiis) .
See also references under yenus.
hasalis Sloane 1907, Deutsche Eut. Zeitschrift for
1907, p. 182 (new synonymy).
Description. None required here; length
c. 6-8 mm.
Types. Of nipicoUis, from Port Denison
( Bowen ) , Queensland, Australia, presum-
ably in Macleay Mus., Sydney; of basalis,
from the Gazelle Pen., New Britain, should
be in Deutsche Ent. Institut, Berlin-
Dahlem ( none seen ) .
Occurrence in New Guinea. Thirty-four
specimens, from localities covering almost
the whole length of New Guinea, most at
low altitudes but records up to 1300 (at
Wau), 1400, and 1500 m.
Notes. Sloane distinguished hasalis from
niii,rieollis by a slight color difference,
which does not hold in the series before me:
5 specimens from New Britain include both
individuals with base of elytra entirely dark
{hasalis) and individuals with the pale
marks reaching the elytral base (nigricollis).
The series from New Ckiinea is even more
variable, with elytra ranging from almost
wholly reddish testaceous to almost wholly
piceous with subbasal pale marks scarcely
indicated. The pronotum also varies, from
reddish piceous with pale margins to red-
dish testaceous, and the variation is partly
independent of the variation of elytral
pattern. All these color forms seem to me
to be one species. The variation of pattern
may prove to be partly geograjihic, but the
material liefore me is not suHicient to
establish this.
Phloeocarobus euplenes n. sp.
Descrij)tion. With characters of genus;
lonn as in T^igure 110; strikingly bicolored,
head and prothorax red, elytra blue, lower
surface and appendages reddish testaceous;
moderateh' shining, reticulate niierosculp-
ture absent or hunt on bout and on disc
oi pronotum, distinct and slightK trans-
verse on cKlra. Head: O.Tf) and 0.76 w idlh
prothorax; front scarcely impressed, faintly
sparsely punctulate. Prothorax: width/
length 1.58 and 1.62; base/apex 1.67 and
1.62; margins broadly flattened, not much
reflexed; disc sparsely irregularly punctu-
late and faintly transversely strigulose.
Elytra: width elytra prothorax 1.42 and
1.47; striae well impressed, faintly punctu-
late; intervals slightly convex, each with an
irregular row of punctules along middle.
Secondary sexual characters: i as for genus;
9 unknown. Measurements: length 5.0-
5.2; width 2.1-2.2 mm.
Types. Holotype 6 (Bishop Mus.) from
Torricelli Mts., Wantipi Village, N-E. N. G.,
Nov. 30-Dec. 8, 1958 (W. W. Brandt);
and 1 i paratvpe (M.C.Z., Type No.
31,490) from Kiunga, Fly R., Papua, T'llv
11-14, 1957 (W. W. Brandt).
Notes. This new Phloeocarahus is strik-
ingly different in color from any other
member of the genus known to me. In
form and color it resembles and may mimic
New Guinean species of the agonine genus
Eui)lenes.
Genus TRIGONOTHOPS Macleay
Macleay 1804, Trans. Eut. Soc. New South Wales
1, p. 110.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1488 (see for additional references and list
of species).
DiaiS.nosis. Sec^ Key to Genera of Lehii)}i
of New Guinea and Notes under the follow-
ing species.
Description. None rt>quired here.
Type s))ecies. Calleida pacifica Erichson
of Australia (original designation).
Generic distribution. TT(>retofore known
onK' horn Australia including Tasmania
and oilier elose-King islands; range now
extended lo New (>uinea.
Notes. The Australian numbers ot this
geinis li\(" on tree trunks. 1 think.
Trigonothops lateralis n. sp.
Description, i-'onii as in f'igure 111; en-
tireU (slightly reddish) xellow except h:)r
a wide biowiiish black stripe along the
The Carabid Beetles of New Guinea
Darlinp,ton
185
outer side of each elytron (not including
the reflexed margin) from humerus nearly
to apex; not pubescent; moderately shining,
with lightly impressed reticulate micro-
sculpture isodiametric on front, slightly
transverse on disc of pronotum and elytra.
Head 0.86 width prothorax; 2 setae over
each eye; front slightly impressed at middle,
longitudinally impressed each side anteri-
orly; clypeus broadly rounded anteriorly
( a small notch at middle may be abnormal ),
1-setose each side; labrum transverse, ir-
regularly broadly rounded in front, 6-setose;
mentum with triangular tooth; ligula thick-
ened, blunt, probably originally setose but
setae broken short; paraglossae c. long as
or slightly shorter than ligula, attached
except at extreme apex, apparently with-
out setae; maxillary palpi not or not much
thickened, labial palpi with apical seg-
ments wider, not quite ^2 wide as long,
narrowly obliquely truncate. Prothorax
subcordate but with broad base; width/
length 1.44; base/apex 1.54; base broadly
briefly lobed, margined; apex broadly emar-
ginate, not distinctly margined at middle;
side margins broad and reflexed especially
toward base, each with 2 seta-bearing punc-
tures, at basal angle and c. % from apex;
disc with middle line deep, transverse im-
pressions less well defined, baso-lateral
foveae deep but not sharply limited; surface
of disc with faint weak transverse strigula-
tion, almost impunctate. Elytra: width
elytra/prothorax 1.94; humeri slightly nar-
rowed and rounded but not obliterated;
reflexed lateral margins moderate; apices
slightly obliquely truncate and very slightly
sinuate, with outer angles broadly and su-
tural angles narrowly rounded; striae entire,
moderately impressed, faintly irregular but
not distinctly punctulate; intervals slightly
convex, not distinctly punctate except 3rd
with 2 small dorsal punctures, on inner edge
just before middle and behind apical Vi (sub-
basal puncture, if present, minute and not
surely detectable). Inner wing,s full. Legs:
hind tarsi missing; middle tarsi with 4th
segments very deeply emarginate, with
lobes much longer than V2 length of seg-
ment; claws (of front tarsi) each with 4
long teeth and apparently an additional
very short tooth toward base; 5th segments
(of front tarsi) with accessory setae. Sec-
ondary sexual characters: i unknown; $
with last ventral segment subtruncate,
slightly subsinuate at middle, with 2 setae
near apex each side. Measurements: length
6.7; width 2.9 mm.
Type. Holotype 9 (Leiden Mus.) from
Wissel Lakes, central West N. G., Arabu
Camp, 1800 m, Oct. 12, 1939 (H. Boschma);
the type is unique.
Notes. Even without the 6 I am rea-
sonably sure that this insect is a Trigono-
thops. It agrees with T. pacificus Erichson
in form and in significant characters in-
cluding the mouthparts (see preceding
Description ) and position of the dorsal
elytral punctures. Moreover, the elytral
color pattern is derivable from that of
Trigonotliops pacificus: lateral stripes like
those of hiteralis would be left if the inner
portion of the dark elytral pattern of
pacificus were erased.
This individual had lost most of its legs
before I received it. Only the left front leg
is still complete, and the left middle leg is
complete except for the 5th segment. But
these are enough to show essential char-
acters, and the specimen is in good condi-
tion otherwise.
Genus NOTOTARUS Chaudoir
Chaudoir 1875, Bull. Soc. Nat. Moscow 49, Part
2, p. 19.
Sloane 1898, Proc. Linnean Soc. New South
Wales 23, p. 494.
Diagnosis. As Aiu)motarus (following
genus) but side pieces of metasternum short,
scarcely longer than wide; and (in the New
Guinean species) genae short-setulose; an-
tennae with segments 2 and 3 more or less
pubescent; tarsi pubescent (sparsely short-
pilose) above; i middle tibiae not modi-
fied.
186 Bulletin Mit.'icum of Coinparatwv Zoolof^y, Vol. 137, No. 1
Description. See Notes (below) and de-
tailed Description of following species.
Ti/j)e species. Nototariis australis Chau-
doir, of Western Australia.
Generic distribution. Previously known
only from Australia; range now extended
to New Guinea.
Notes. Chaudoir described Nototarus as
without a mentum tooth, and the tooth is
certainly difficult to see in some Australian
species, but it may be depigmented rather
than absent. The characters and generic
classification of this group of Carabidae
need further study, which will have to be
based on the Australian rather than New
Guinean forms. Some Australian species
assigned to Nototarus by Sloane do have
the mentum toothed, and the single New
Guinean species (below) is evidently
closely related to some of them.
Nototarus papua n. sp.
Description. Form as in Figure 112;
brownish black, humeri broadly paler
brown, appendages brownish testaceous;
reticulate microsculpture lightly impressed,
c. isodiametric on front, slightly transverse
on pronotum, more irregular on elytra, and
surface irregularly rather sparsely punctu-
late. Head 0.81 and 0.80 width prothorax;
eyes moderately prominent, genae rounded-
oblique, short-setulose; 2 setae over each
eye; front longitudinally rugulose each side;
clypeus slightly emarginate-truncate, 1-
setose each side; labrum wide, slightly
emarginate-truncate, 6-setose; mandibles
rather short, curved; antennae moderate,
pubescent from middle of 3rd segments;
mentum w ith long, narrowly rounded tooth;
ligula 2-setose, paraglossae attached, e(iual
in length, wide, not setose; maxillary palpi
slender, labial palpi with apical segments
wide. Prothorax cordate, short-lobed at
base; width length 1.31 and 1.30; base
apex 0.88 and 0.92; margins narrow, each
with seta at basal angle and c. ' i from apex;
basal and apical marginal lines inti-rrupted
at middle; disc with middle line coarse,
almost entire, transxcrse impressions almost
obsolete. Elytra short, narrowed toward
base, connate; width elytra/prothorax 1.49
and 1.51; striae entire, impressed, faintly
punctulate; intervals convex especially
toward base, 3rd 2-punctate, the punctures
near middle of length and c. V4 from apex.
Inner icings vestigial. Lower swjace not
obviously pubescent ( in part with very
short inconspicuous sparse pubescence).
Le^s moderate; tarsi short-pilose above;
4th segments middle and hind tarsi slighth"
emarginate; 5th segments with long ac-
cessory setae; claws each with 3 or 4 teeth.
Secondary sexual characters: labial palpi
with apical segments wider in c^ (truncate
apex wider than length of inner edge),
narrower in 9 (truncate apex narrower
than inner edge); 6 front tarsi slightly
( scarcely ) dilated, 3 segments 2-seriately
squamulose; i middle tarsi not squamu-
lose; i middle tibiae not modified; $ with
1, 92 seta-bearing punctures each side
last ventral segment. Measurements: length
4.6-5.3; width 1.9-2.3 mm.
Types. Holotype 6 (M.C.Z., Type No.
31,491) and 20 paratypes all from Dobo-
dura, Pa|>ua, Mar.-July 1944 (Darlington).
Measured specimens. The c^ holot\'pe and
1 9 paratype.
Notes. This is the onl\' Nototarus thus
far found in New Guinea. It may be re-
lated to N. morosus Sloane of Port Darwin,
Australia, but has thc^ prothorax exidently
more narrow(>d posteriori), with sinuate
sides. Other, apparently related, unde-
scribed spicies occur in North Queensland.
This species is one of the very few strictly
flightless Garabidae found at low altitudes
in New Guinea. Most or all of in\- speci-
mens were, I think, taken in flood debris
from the floor of rain forest. The rain-forest-
floor habitat and the insect's tlightlessness
pi'rhai)s explain w h\ other collectors have
not tound it.
Genus ANOMOTARUS Chaudoir
Chaiuloii I,S7.^, linll. Soc. Nat. Moscow U), Part 2,
p. \H.
The Carabid Beetles of New Guinea • Darlington 187
Sloane 1898, Pioc. Linnean Soc. New South Wales
23, p. 494.
Csiki 1932, Coleop. Cat., Caraliidae, Harpalinae 7,
p. 1493 ( see for additional references, synonymy,
and list of species ) .
Tedlicka 1963, Ent. Abhandlungen 28, pp. 300,
450.
Diap,nosis. See Key to Genera of Lebiini
of New Guinea.
Deseription (based on New Giiinean
species only). Fomi as in Figures 113-
117; small, slender, depressed; elytra usu-
ally (not always) with characteristic pale
marks; not pubescent. Head: eyes mod-
erately prominent, genae rounded, more or
less prominent ( usually less so than eyes ) ;
2 setae over each eye; antennae moderate,
pubescent from 4th segment (3rd segment
with only usual apical setae); clypeus trans-
verse with rounded angles, 1-setose each
side; labrum transverse, broadly emargi-
nate, 6-setose; mandibles short, strongly
curved; mentum with long tooth; ligula
rather broad, 2-setose, paraglossae attached
and c. equal in length, wide, not setose;
maxillary palpi moderate, not widened;
labial palpi with apical segments widened.
Frothorax cordate, with base briefly lobed;
side margins moderate or narrow, with
setae at basal angle and c. V-i or % from
apex; disc with impressed middle line and
less distinct transverse impressions. Elytra
with apices simple; striae entire, not dis-
tinctly punctate; 3rd intervals 2-punctate,
with punctures near or before middle and
c. Vi from apex; 8th intervals usually finely
carinate on inner edge near base (carinae
sometimes so fine as to be scarcely detect-
able). Inner wings iuW. Lotver swiace not
extensively pubescent; side pieces of meta-
sternum long. Legs slender; tarsi not pubes-
cent above; 4th segments middle and hind
tarsi slightly emarginate; 5th segment with
long accessory setae; claws with c. 4 or 5
teeth. Secondary sexual characters: i front
tarsi slightly (scarcely) dilated, 3 segments
2-seriately squamulose; last segment i labial
palpus wider (truncate outer edge almost
as long as inner side), of 9 less wide; 6
middle tarsi without squamae; i middle
tibiae with inner edge tuberculate-serrate
( cf . Demetrida ) , with c. 3 or 4 low tuber-
cles in row toward apex ( in all New
Guinean species of which the c5 is known ) ;
i with 1, 9 2 setae each side last ventral
segment.
Type species. Anomotarus olivaceus
Chaudoir, from Melbourne, Australia.
Generic distribution. Southern Asia
(Ceylon, India, Japan, etc.) to Australia
and Tasmania, and New Caledonia.
Notes. Although I recognize 8 (closely
interallied) species of Anomotarus in New
Guinea, material is scanty and almost
nothing is known of their habits. I think
most of them probably live among dead
leaves on the ground in rain forest. A few
specimens have been taken at light.
Key to Species of Anomotarus of New CIuinea
1. Each elytron either with longitudinal post-
humeral stripe (outside 4th stria) and sub-
apical-sutural spot pale, or with only the
subapical spot, or unmarked 2
- Each eh tron with a more or less incomplete
oblique or transverse (not longitudinal) spot
or band before middle and usually (not
always ) a subapical-sutural spot pale 5
2. Very slender (prothoracic width/length
1.15); subsericeous black, unmarked (p.
188 ) gressitti
- Less slender (prothoracic width/length c.
1.20 or more); elytra with at least subapical-
sutural pale spot(s) 3
3. Elytra with distinct post-humeral stripes
(as well as subapical-sutural spot(s)) pale;
prothorax less narrowed basally (base/apex
e. 1.10 or more) (p. 188) (stignnila)
- Post-humeral stripes indistinct or absent
(subapical-sutural spot(s) distinct); pro-
thorax more narrowed basally (base/apex
1.05 or less) 4
4. Brown, with subapical-sutural spots small and
separated; prothorax narrowed (width length
1.22 and 1.23); elytra more narrowed toward
base (p. 188) —_ tvallacei
- Black, with subapical-sutural spots united in
conspicuous square plagia; prothorax wider
(width length 1.35); elytra less narrowed
toward base ( Eig. 115) (p. 189) phigifer
5. Each elytron with transverse-oval pale spot
before middle but without subapical spot
(Eig. 114) (p. 189) ocellatus
188 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
- Elytra with aiitt'iior and posterior pale
spots 6
6. Elytra with an almost strictly transverse,
rej^ular, c. entire pale fascia before middle
( as well as an incomplete siibapical fascia ) ;
elytral margins bicolored; femora bicolored;
surface in part sericeous; length 5.5-6.0 mm
(p. 189) tratisversus
- Anterior elytral marks more oblicjue and/or
more irregular and/or more interrupted;
elytral margin usually entirely translucent-
testaceous; femora not bicolored; surface not
or not so strongly sericeous, more shining;
size usually smaller 7
7. Anterior elytral marks usually more oblique;
femora pale ( Moluccas and western antl
central New Guinea) (p. 190) onuitii.s
- Anterior elytral marks more nearly trans-
verse; femora darker (central and eastern
New Guinea) 8
8. Prothorax wider (width/length 1.38 and
1.41); markings wide (p. 190) fuscipcs
- Prothorax narrower (width/length 1.27);
markings n;irrower (Fig. 117) (see also
Dc'scii)>tiou) (p. 191) uaii
Anomofarus gressitti n. sp.
Description. With cliaractcrs of geiui.s;
form as in Figure 113; .slender; lilack, not
marked, but margins of prothorax and espe-
eially of elytra more or less pale translucent;
surface dull, closely punctulate and micro-
reticulate, elytra subalutaceous; legs testa-
ceous, antennae and mouthparts brownish
testaceous. Head 0.80 width prothorax.
Frothorax: width length 1.15; base, apex
1.00. Elytra: width elytra prothorax 1.73;
striae lightly iinpressed, faintly punctulate;
intervals c. flat, 6th, 7th, and (Sth with
inner edges finely carinate tor increasing
distances from base. Measiireitient.s: length
c. 5.0; width 1.9 mm.
Type. Holotype 9 (Bishop Mus.), from
Maprik, N-E. N. G., 160 m, Oct. 14, 1957
(Gressitt); the type is uniciue.
Notes. The slender form and dull bkuk
color, without markings, should nuikc this
species easy to recogni/e.
(Anomofarus stigmula (Chaudoir))
Ghaudoir 18.52, Bull. Soc. Nat. Moscow 25. Part 1,
p. 57 {Ci/min(Jis) .
Andrewes 19.30, C^at. Indian insects 18, (^arabidac,
p. 28.
Csiki 1932, Goleop. Gat., Garabidae, Ilarpalinae 7,
p. 1493 (see for synonymy and additional refer-
ences ) .
Louwerens 1953, Verhandlungen Naturforschcnden
Gesellschaft Basel 64, p. 316.
Jedlicka 1963, Ent. Abhandlungen 28, p. 451.
Description. With characters of genus;
form, including elytra, elongate; brown,
elytra each with longitudinal humeral mark
( outside 4th stria ) and variable apical mark
pale; appendages including femora testa-
ceous; rather shining, but surface finely
microreticulate and sparsely inconspicu-
ously punctulate. Head 0.84 and 0.86 width
prothorax. Prothorax: width length 1.24
and 1.29; base /apex 1.14 and 1.11; margins
moderate, with basal angles well defined, c.
right or obtuse (not acute). Elytra long;
width elytra prothorax 1.65 and 1.69; striae
moderately impressed, not distinctly punc-
tulate. Mea.suremcnts: length c. 4.7-5.3;
width c. 1.9-2.1 mm.
Type. From Simla(h), northern India;
now in Oberthiir Coll., Paris Mus. (not
seen ) .
Occurrence in New Guinea. Doubtful;
see following Notes.
Measured specimens. A i from Coim-
batore. South India, and 9 from Lawa,
Malita, Davao Prov., Mindanao, Philippine
Islands (both specimens M.C'.Z. ).
Notes. This species is now recorded
over a very wide area, from SE. Asia
including Ceylon and Japan to Timor and
New Caledonia, but apiiarentb not .\us-
tralia. New Cluinea is included in the
species' range by Andrewes and Csiki, but
I have found no detailed record of its oc-
currence there. I suspect its supposed oc-
cuiicnce is erroneous, b;ised on the old
specimens in the British Museum described
below ;is ualhuci. I h;i\(- not seen true
sliii,ninla Irom New (iuiiiea.
Anomofarus wallacei n. sp.
Dcscripliou. With c h;ir;icteis ol genus;
lonn shorter than usual in genus; brownish
pieeous, el\'tr;i e;uli willi laint p;iler AVCd
behind Immeius (corresponding to post-
The Carabid Beetles of New Guinea
Darlington
189
humeral spot of stigmula), a small testa-
ceous spot on intervals 2 and 3 just before
apex, and lateral margin narrowly brownish
testaceous; appendages brownish testa-
ceous; surface (in part) moderately shining,
with reticulate microsculpture lightly im-
pressed and punctulation rather sparse.
Head 0.84 and 0.89 width prothorax. Pro-
thorax: width/length 1.23 and 1.22; base/
apex 1.00 and 1.05. Elytra shorter and more
narrowed basally than usual in genus; width
elytra/prothorax 1.69 and 1.74; striae mod-
erately impressed, not distinctly punctulate.
Inner wings apparently fully developed in
spite of narrowing of humeri. Measure-
ments: length 4.3-4.9; width 1.8-2.0 mm.
Types. Holotype 5 (British Mus.) and
1 9 paratype (M.C.Z., Type No. 31,492)
both from Dory, West N. G. (presumably
collected by Wallace); c^ unknown.
Notes. This species is very close to
stigmuhi (above) but has elytra shorter,
more narrowed at base, and with markings
reduced, and the prothorax coaptively nar-
rowed at base, as the ratio base apex shows.
It may prove to be a geographic subspecies
of stigmida. It may prove not to be from
New Guinea (because the "Dory" locality
is always dubious), but it seems not to be
known anywhere else.
Anomofarus plagifer n. sp.
Description. With characters of genus;
form as in Figure 115; brownish black,
elytra with conspicuous, common, square
spot just before apex pale and lateral
margins slightly pale translucent; append-
ages testaceous; surface rather shining but
lightly microreticulate and faintly sparsely
punctulate. Head 0.88 width prothorax.
Prothorax strongly cordate (more so than
in stigmida); width/length 1.35; base/apex
1.02; posterior angles abruptly right-acute.
Elytra of moderate length, slightly nar-
rowed toward base ( less than in walktcei ) ;
width elytra prothorax 1.57; striae well im-
pressed, not distinctly punctulate. Measure-
ments: length c. 4.9; width c. 2.0 mm.
Type. Flolotype S (Bishop Mus.) from
Port Moresby, Papua, May 20, 1956
(Gressitt), taken in light trap; the type is
unique.
Notes. As compared with stigmida,
plagifer is slightly broader, with more
cordate prothorax, and is darker in color,
without basal but with more conspicuous
subapical elytral marks.
Anomofarus ocellafus n. sp.
Description. With characters of genus;
form as in Figure 114; black (bluish black
in some lights), each elytron with trans-
verse-oval pale spot before middle between
striae 1 and 7; appendages brownish testa-
ceous, femora darker brown; shining, reticu-
late microsculpture light ( faint on part of
pronotal disc) and punctulation very fine,
faint, sparse. Head 0.85 width prothorax.
Prothorax cordate; width/length 1.31; base/
apex 0.96; side margins narrower than
usual. Elytra: width elytra prothorax 1.59;
striae moderately impressed. Measure-
ments: length 4.4; width 1.8 mm.
Type. Holotype 9 (Louwerens Coll.,
eventually to Leiden Mus.) from Sorong
"Kpg. Roefci," West N. G., July 8-Aug. 14,
1948 (M. A. Lieftinck); the type is unique.
Notes. The small size, relatively shining
surface, and unique markings distinguish
this species.
Anomofarus fransversus n. sp.
Description. With characters of genus;
form as in Figiue 116; large, with wide-
cordate prothorax; dull aeneous black,
elytra more alutaceous, with c. entire cross-
band before middle and incomplete trans-
verse mark before apex pale; elytral margins
bicolored, pale at transverse fascia, dusky
elsewhere; femora bicolored, dark with
pale apices; appendages otherwise brown-
ish testaceous, antennae slightly darker
distally; surface closely microreticulate,
sparsely punctulate. Head 0.72 and 0.76
width prothorax. Prothorax: width length
1.46 and 1.43; base apex 1.08 and 1.03;
posterior angles abruptly acute. Elytra:
190 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
width elytra/prothorax 1.46 and 1.44; striae
fine, lightly impressed, not punctulate; in-
tervals almost flat. Measurements: length
c. 5.5-6.0; width 2.1-2.3 mm.
Types. Holotype S (M.C.Z., Type No.
31,493) and 1 9 paratype both from Do-
bodura, Papua, Mar.-July 1944 (Darling-
ton); 3 paratypes, Popondetta, Papua, 60
m, Aug. 30-31, Sept. 1-4, 1963 (Sedlacek),
light trap.
Notes. The large size, dull partly aluta-
eeous surface, and color pattern, especially
the evenly transverse anterior elytral fascia,
characterize this species.
Anomofarus ornafus Louwerens
Louwerens 1956, Treubia 23, p. 237.
Description. With characters of genus;
form c. as of following species (fuscipcs);
piceous black, elytra each with broad pale
mark before middle, this mark being ir-
regularly transverse-oblique with outer-an-
terior corner extended toward and some-
times reaching humerus, and elytra also
with subapical-sutural spot pale and lateral
margins entirely pale; legs entirely testa-
ceous; antennae brownish testaceous; sur-
face shining, reticulate microsculpture light
on head, faint on disc of pronotum, more
distinct on elytra. Head 0.cS6, 0.(S5, and
0.83 width prothorax. ProtJiora.x cordate;
width length 1.43, 1.36, and 1.33; base/apex
1.08, 1.07, and 1.08; posterior angles
abruptly right or acute. Elytra: width
elytra prothorax 1.72, 1.77, and 1.77;
striae moderately impressed, not punctu-
late. Measurements: length 5.1-5.8; width
2.2-2.4 mm.
Types. From Obi Is. (Laiwui, 0-200 m.
Sept.-Oct. 1953, A. M. R. Wegner); holo-
type in Leiden Mus. (not seen).
Occurrence in Neic Guinea. West N. {,.:
8, Cyclops Mts., Sabron, Camp 2, 2000 ft.
(610 m), July 1936 (Cheesman); 4, vie.
Ilollandia (various collectors); 1, Kebar
Vy. W. of Manokwari, 550 m, Jan. 4-31,
1962 (S. & L. Quale, Bishop Mus.), in light
trap; 1, Sansapor, Aug. 1944 (Darlington).
Measured .specimens. A pair (69) from
Cyclops Mts. and a 9 paratype from Obi
Is.; figures given in this order.
Notes. See under the following species
( fuscipes ) .
Anomofarus fuscipes n. sp.
Description. With characters of genus;
form average; black ( slightly brownish or
aeneous), elytra each with slightly oblique
transverse fascia just before middle and
subapical-sutural spot pale and lateral
margins entirely pale brownish translucent;
femora brown (not distinctly bicolored),
tibiae and tarsi paler; antennae brown;
upper surface rather shining, reticulate
microsculpture light especially on disc of
pronotum. Head 0.76 and 0.77 width pro-
thorax. Prothorax cordate; width length
1.38 and 1.41; base apex 1.17 and 1.09;
basal angles abrupth' right or acute.
Elytra: width elytra prothorax 1.66 and
1.59; striae moderately impressed, not punc-
tulate. Measurements: length 4.1-5.5;
width 1.9-2.3 mm.
Types. Holotype $ (M.C.Z., Type No.
31,494) from Dobodura, Papua, Mar.-July
1944 (Darlington); and paratypes as fol-
lows. Papua: 1, Karema, Brown R., Mar.
8-11, 1955 (E. O. Wilson, M.C.Z.), taken
in lowland rain forest; 2, Ml. Riu, Sudest
Is., 250-350 m, "No. 10," Sept. 3, 5, 1956
(L. J. Brass, A.M.N.H.); 1, Abaleti, Rossel
Is., 0-50 m, "No. 12," Oct. 9, 1956 (Brass,
A.M.N.H.). N-E. N. G.: 1, vie. Nadzab,
Julv 1944 (Darlington); 1, same localitv,
May 20-22, 1955 (E. O. \Ailson, M.C.Z.'),
in dry e\'ergr{>en forest; 1, Eriina, Astrolabe
Bav, 1896 (Biro); 1, Stephansort, Astrolabe
Ba'v, 1898 (Bin')). West N. G.: 4, Ilol-
landia, Nov. 21, 1944; Ma\ 1945; May 4,
1947 (Iloogstraal, M.C.Z. ); 1, same localitv,
May 1945 ( Malkin, U.S.N.M.).
Measured si)ecinu'ns. The • holot\pe and
1 9 parat\pe from Ilollandia.
Notes. This species is very close to the
[M'eeeding (ornatus), but is distinguished
by brown lather than testaceous femoia
and 1)\ more transverse anterior eKtral
The Carabid Beetles of New Guinea • Darlinaton 191
fascia which approaches the humeri less
closely, although this mark varies somewhat
in both species. A. fuscipcs occupies ap-
proximately the eastern half of New Guinea,
oniaftis the western half and the Moluccas,
but their ranges are not strictly allopatric,
since both occur in the vicinity of Hol-
landia.
Anomotorus wau n. sp.
Description. With characters of genus;
form as in Figure 117; black, elytra each
with oblique fascia before middle and with
common subapical-sutural spot testaceous
and margins brownish translucent; append-
ages brownish testaceous with femora
darker; shining, reticulate microsculpture
and fine sparse punctulation present but
light especially on pronotum. Head 0.84
width prothorax. ProtJiorax narrow-cordate,
with relatively narrow margins; width
length 1.27; base apex 1.05; sides more
oblique posteriorly (less broadly rounded)
than in fmcipes, with posterior angles
abruptly e. right. Elytra: width elytra
prothorax 1.67; striae well impressed, not
punctulate. Measurements: length 5.1;
width 2.0 mm.
Type. Holotype 9 (Bishop Mus.) from
Wau, Morobe Dist., N-E. N. G., 1200 m,
July 5, 1961 (Sedlaceks), taken in light
trap; the type is unique.
Notes. Although this may be only a form
of the preceding species (fiiscipes), it has
a relatively narrow prothorax and reduced
elytral fasciae and will probably prove to
be worth distinguishing. More material
from more localities is needed to show
whether it is a species or a geographic
subspecies.
Tribe PENTAGONICINI
Fentugonicidac Auct. including Jeannel 1949,
Coleop. Carabiques de la Region Malgache,
Part 3, p. 767.
Jeannel 1942, Faune de Fianee, Coleop. Carabiques,
Part 2, p. 1017, footnote.
Penta^onicinae Basilewsky 1953, Exploration Pare
National TUpemba, Fasc. 10, p. 183.
Scopodmi Csiki 1932, Coleop. Cat., Carabidae,
Harpalinae 7, p. 1500 (see for synonymy and
additional references).
Jedlicka 1963, Ent. Abhandlungen 28, p. 505.
The beetles of this tribe resemble Lebiini
but (according to Jeannel 1949) are not
related to them. Pentagonicini can usually
be recognized at a glance by form, and
the tribe is defined by technical characters
including obliteration of the suture that,
in most Carabidae, separates the mentum
from the base of the head posteriorly.
However, this suture is still indicated in
Parascopodes.
This tribe consists of four genera. One,
Pentagonica., occurs in all the warm regions
of the world. One, Parascopodes (de-
scribed below), consists of a single species
that occurs in both eastern New Guinea
and northeastern Australia. One, Actenonyx
(1 species), is confined to New Zealand.
And the fourth genus, Scopodes, is best
represented in Australia and extends to
New Zealand and to mountains on New
Guinea and on Java. Ecologically, Pen-
tagonica alone of these 4 genera is pri-
marily arboreal, occurring especially in
masses of vines and other vegetation near
the ground, although some species are
found among dead leaves on the ground.
Parascopodes cyaneus occurs, in my limited
experience, in grass or on the ground under
grass (I am not sure which). The New
Zealand genus (Actenonyx) is probably
ground-living. And Scopodes is ground-
living but some species occur on logs or
tree trunks.
The distribution of genera of Penta-
gonicini suggests two possible geographic
histories. The tribe may once have been
better represented in other parts of the
world and may have withdrawn (or may
be withdrawing) into the Australian Region.
Or the tribe may have originated in Aus-
tralia and diversified there, and Pentagonica
may have spread from there over the rest
of the world, its spread perhaps facilitated
by its invasion of arboreal habitats in
which flight and dispersal may have been
192 BiiUctin Miiscinu of Comparative Zoology, Vol. 137, No. 1
favored. The very wide distributions of
some species of Pento^onica show that the
insects do disperse readily. Of course, there
is a third possil^ihty, that the geographic
liistory of the tribe has been more complex
than can be guessed from present distribu-
tions of genera and cannot now be deci-
phered at all. Nevertheless, the history of
the tribe is worth guessing about. Its
distribution may become more significant
if other Carabidae or other animals are
found to have similar geographic patterns.
There is, for example, a suggestive general
similarity between the distributions of this
tribe of carabid beetles and of the parrots,
which are most diverse in the Australian
Region with one of the several Australian
subfamilies spread over the warmer parts
of the world (Darlington, Zooiieoiiraplnj,
Wiley, 1957, pp. 271-272, ;30()-;301, fig. .34).
Key to Genera of Pentagoxicim of
New Guinea
1. Form Lt'/;;'a-like, with eyes only iioniuilly
prominent (Fig. 118) (p. 192) _.._ Pcntd^onica
— Fomi more eompaet, with eyes larger and
more abruptly pronnnent (Figs. 120, 121) _ 2
2. Ligula normal, not miieh swollen, mueli
shorter than paraglossae; i front tarsi 2-
seriately squanmlose; elytral striae and in-
tervals regular, without eonspieuous foveae
(p. 19.5) Parascopodes
- Ligula swollen, club-like, as long as or longer
than paraglossae; i front tarsi with soles of
densely packed slender squamae; elytral
striae and intervals usually more or less ir-
regular, .^rd intervals usually with eon-
sj^iiciinns lovcae (p. 197) _ Scopixlcs
Genus PENTAGONICA Schmidt-Goebel
Schmidt-( ioclx'l I S4(), I'auniila (]olco|). Hiiiiianiae,
p. 47.
Csiki 19.32, Clolcop. Gat., ("arahidae, llarpaiiiiac 7,
\i. 1.500 ( see lor additional r(i<icnees, s\iion\niy,
and list of species ).
Di(iu.n()si.s. Immediately recogni/able b\-
lorm ( I'ig. 11<S) and tribal characters.
l)c.scrij)liou. None reciuired here.
Tijjx' species. Peiitagonica ntficollis
Schmidt-Cioebel ( below ) .
Generic distribution. All warm regions
ol the world.
Notes. The members of this genus are
winged, active, and apparently diurnal.
They usually live in dense vegetation within
a few feet of the ground, or sometimes in
leaf litter on the ground. Specific char-
acters in the genus arv few, principally
slight differences of form, microsculpture,
and color pattern. There is some individual
variation, including apparent dimorphism
of color in some cases. And understanding
of the species is made more difficult by the
very wide distributions of some of them.
Because the prothorax has no distinct an-
terior angles, the ratio base apex is omitted
in the following specific descriptions.
Key to Species of Pentagonica ok New Guinea
1. Prothorax strongly pedunculate; ( bicolored,
dark with prothorax red; elytral striae punc-
tulate; size small, length .3-.3.8 mm) (p.
192) pallipcs
- Prothorax less strongly pedunculate; size usu-
ally larger 2
2. Pronotum with lateral margins usually con-
nected posteriorly by a weak transverse
ridge that is not ((uite basal; reticulate
microsculpture of elytra often somewhat
transverse; (color above unitormK d;irk or
with prothorax red; length c. 3.5-4.5 mm)
(p. 193) l^himla
- Proncjtum with lateral margins connected
posteriorly b>' a line ridge that reaches
extreme base; elytral microreticulation usu-
ally c. isodiametric; size larger .3
3. Prothorax relatively narrower ( width length
1.57 and 1.65); (color dark, with lateral
margins of prothorax and elytra strikingh'
pale) (p. 194) crichsoui
- Prothorax relatively wider (width length
1.72-1.79) [ 4
4. Bicolored, d;uk with red prothorax (p.
194) nificollis
- Not bicolored, entirely dark 5
5. Flytral striae clearly indicated (hut scarcely
impressed); antennae rilativeb dark ;uid
femora relatixcK pale (p. 194) papiia
- FKtral striae \irtu:dl\ obsolete; antennae
testaceous, Icmoia rclatixcK (!;nk (p. 19.5)
_ cstriaia
Pentagonica pallipes (Nietner)
Nielncr 18.5(i, 1. Asiatic Soc. Bengal 25, p. .525
(Klliolia).
(.'siki 1932, (^oK'op. i'.ni.. Gar:ibida<', Ihupalinae 7,
p. 1.502 (see for sNiionvinx and additional refer-
ences ).
The Carabid Beetles of New Guinea • Darlington
193
Jedlicka 1963, Ent. Abliandlungen 28, pp. 505,
507.
Description ( for recognition only ) . Head
and elytra dark, prothorax red. Head 0.91
and 0.93 width prothorax. Prothorax pedun-
culate; width length 1.71 and 1.59. Elytra:
width elytra/ prothorax 1.62 and 1.79; striae
punctate; reticulate microsculpture slightly
irregular but scarcely transverse. Measure-
ments: length 3.0-3.8; width c. 1.5-1.9 mm.
Type. From Ceylon; should be in Stettin
Mus. (not seen).
Occurrence in Neiv Guinea. Nineteen
specimens from all 3 political divisions of
New Guinea; most from low altitudes (in-
cluding Oro Bay near Dobodura) but 2 at
1200 m at Wau. One specimen ( from Wau )
is labeled as taken at light.
Measured specimens. A 6 from Hollandia
and 9 from Port Moresby.
Notes. The known range of this rela-
tively distinct species is from Ceylon, the
Malay Pen., etc., to the Philippines, New
Guinea, New Britain, and mid-peninsular
Cape York, Australia (collected hv me
in 1958).
Pentagonica blanda Andrewes
Andrewes 1929, Tijdschrift voor Ent. 72, pp. 315,
339.
?htzocn.'iis Jedlicka 1934, Sbornik Ent. Mus. Prague
12, p. 123.
?})hili]>pmeims Jedlicka 19.34, Sbornik Ent. Mus.
Prague 12, p. 124.
?hottclwri Jedlicka 1935, Acta Soc. Ent. Czecho-
slovakia 32, p. 140.
?eurijodes Andrewes 1938, Ann. Mag. Nat. Hist.
(l"l) l,p. 207.
?quadratipennis Louwerens 1956, Treubia 23, p.
236.
Description ( for recognition only ) . Form
(Fig. 118) broad; color above eitlier en-
tirely dark or dark with red or reddish
prothorax; antennae brown or testaceous;
legs pale, often with darker femora. Head
0.77 and 0.74 width prothorax. Prothorax:
width/length 1.83 and 1.80; lateral margins
usually connected by a poorly defined
(sometimes vague) transverse prebasal
ridge. Elytra: width elytra prothorax 1.51
and 1.54; sutural angles usually blunted or
narrowly rounded but sometimes denticu-
late; striae lightly impressed, usually finely
punctulate ( but variable ) ; reticulate micro-
sculpture often ± transverse. Measure-
ments: length c. 3.5-4.5; width c. 1.6-1.8
mm.
Types. Of blanda, from Sumatra, in
British Mus. (seen); of Jedlicka's species,
from the Philippines, types of luzoensis
and philippinensis in Jedlicka Coll. (not
seen), of bottcheri, in British Mus. (seen);
of euryodes, from Java, in British Mus.
(seen); of quadratipennis, from Halmahera,
Molueeas, in Leiden Mus. ( not seen ) .
Occurrence in Neic Guinea. Very com-
mon throughout New Guinea: c. 130
specimens (about half from Dobodura),
most at low altitudes but a few up to 2500
m (at 1200 m at Wau).
Measured specimens. A pair (69) from
Dobodura.
Notes. I am not ready to synonymize the
names listed above under blanda, but I
suggest that they may all prove to apply
to a single variable species or to members
of a group of very closely interrelated
species that ranges from SE. Asia across
the Malay Archipelago to NE. Australia.
This species or species group varies strik-
ingly in color pattern (individuals uni-
colored or bicolored) and to some extent
in proportions, size, degree of paleness of
margins and appendages, and distinctness
of punctures of elytral striae. Most in-
dividuals from New Guinea are entirely
dark above, but 3 from Finschhafen, N-E.
N. C, and 8 from Hollandia, West N.
C, have the prothorax red. These super-
ficially resemble ruficollis (second follow-
ing species ) , but have different prothoracic
bases, usually rounded sutural angles, and ±
transverse elytral microsculpture. Except
for the red prothorax, these individuals do
not seem to differ from specimens with
dark prothorax taken at the same localities.
At these localities the species is apparently
dimorphic in color of prothorax. However,
at some other localities intermediates oc-
194 Bulletin Musctmi of Comparative Zoology, Vol. 137, No. 1
cur: 4 examples that I haxe from Sansapor
( Vogelkop), West N. G., have the prothorax
paler than head and elytra but reddish
brown rather than clear red. Obviously
this species or group ot species requires
further study, of material from outside as
well as inside New Guinea, before its varia-
tion can be understood.
Most of the many specimens that I col-
lectt^d in New Guinea were taken by day,
by sweeping low vegetation. However, a
few individuals, including both color forms
at Finschhafen, are from light-trap material
and evidently flew to light at night.
I collected numerous dark (not bicolored)
specimens apparently of this species at
several localities in North Queensland in
1957-195(S. The species seems not to have
received a name in Australia.
Penfagonica erichsoni Schmidt-Goebel
Schmidt-Coel^el 1846, Faunula CoKop. Biniianiat',
p. 48.
Csiki 1932, Coleop. Cat., Carabidat', llaipalinae 7,
p. 1501 (see for synonymy and additional refer-
ences ) .
Jedlieka 1963, Knt. Ahliandlnntjen 28, pp. 506.
.511.
Description ( for recognition only), h^orm
of large Fcntuiionica with rather narrow
prothorax; dull black, reflexed margins of
prothorax and elytra very pale or translu-
cent, legs testaceous, antennae brown.
Head 0.85 and 0.84 width protliorax. Pro-
thorax: width length 1.57 and 1.65; margins
posteriorly connected b\ a fine curved
ridge across extreme base. Elytra: width
elytra/ prothorax 1.82 and 1.78; sutural
angles usually denticulate; striae sHghtl\
impressed, vaguely or not distinctl\ punc-
tate; elytral microreticulation r. isodiamet-
ric. Measurements: length e. 5.0-5.5;
width e. 2.0-2.2 mm.
Type. l<'rom liurma; in Prague Mus. (not
seen ) .
Occurrence in Netc Guinea. Tapua: 1,
Dobodura, Mar.-lul\ 1944 (Darlington);
1, Woodlark Is. (Murn;i), Kulumadau Hill,
Apr. 20-30. 19.57 (W. W . Brandt, l^isliop
Mus.). N-E. N. G.: 1, Simbang, Huon
Gulf, 1898 (Biro); 1, Wau, 1200 m, Sept.
15-30, 1962 (Sedlacek); 1, Wum, Upper
Jimmi Vy., 840 m, July 16, 1955 (Gressitt).
Measured specimens. Two $ 9 , from
Dobodura and Wum.
Notes. P. erichsoni ranges from Ceylon
and SE. Asia to New Guinea and mid-
peninsular Cape York, Australia ( 1 speci-
men. Rocky R., 1958, taken by myself).
Penfagonica ruficollis Schmidt-Goebel
Sehmidt-Coebel 1846, Fannnla Coleop. Birmaniae,
p. 48.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1503.
Jedlieka 1963, Ent. Abhandlunsen 28, p. 509.
Description (for recognition only ). Form
of large Pentagonica; black, prothorax red,
appendages usually black or brown. Head
0.81 and 0.77 width prothorax. Prothorax:
width length 1.79 and 1.78. Elytra: width
elytra prothorax 1.59 and 1.61; sutural
angles usually denticulate; striae very light
or c. absent; microsculpture c. isodiametric.
Measurements: length c. 5.0-5.7; width e.
2.0-2.3 mm.
Type. From Burma, should be in l^rague
Mus. (not seen).
Occurrence in New Guinea. Twt'nt\-six
specimens from Papua, N-E. N. (i.. anil
West N. (i., at low altitudes and up to
1950 m.
M easurrd sjx'cimens. A i")air ( ci 9 ) b-om
Dobodura, Papua.
Notes. The known range of ruficollis is
Irom SE. Asia to .Australia.
Penfagonica papua n. sp.
Description. Form (Fig. 119) of large,
rather slender Pentaiionica: entireK' black
or piecous except suture sonictiuies reddish,
margins sometimes reddish hut not con-
trastinglx pale; h-mora brown, tibiae and
tarsi testaceous; antennae brown with 1st
segments darker biowii; rt'tieulate micro-
seulpture isodiametric on head and pro-
notum. nioic incgnlai- or slightK tiansvcrse
on eKtra. Head 0.78 and 0.78 width pro-
tliorax; labium transxcrse. 6-setose ante-
The Carabid Beetles of New Guinea • Darlington
195
riorl) , the 4 inner setae mueh smaller than
outer ones; mentum without tooth; ligula
not much swollen, 2-setose; paraglossae
slightly shorter than ligula and apparently
attached to it. Prothorax: width length
1.73 and 1.72; margins posteriorly connected
1)\ basal loop. Elytra: width elytra pro-
thorax 1.63 and 1.63; sutural angles usually
narrowly rounded, not denticulate; striae
very finely and lightly indicated, scarcely
impressed, irregularly punctulate; 3rd in-
tervals apparently usually with 3 minute,
well spaced punctures but latter small, dif-
ficult to find, perhaps sometimes absent.
Secondary sexual characters: £ front tarsi
scarcely dilated but with slender squamae
in 2 slightly irregular rows; c^ with 1, 9
2 setae each side last ventral segment.
Measurements: length 4.8-5.7; width 1.8-
2.2 mm.
Types. Holotype $ (M.C.Z., Type No.
31,495) and 10 paratypes all from Dobo-
dura, Papua, Mar.-July 1944 (Darlington).
Measured s})ecimei}s. The 6 holotype and
1 9 paratype.
Notes. The preceding Description gives
all the characters that seem to me worth
mentioning in this genus, in which the
species are so similar to each other. The
particular characters that separate this
species from others in New Guinea are
indicated in the Key to Species. The present
new species is larger than blandii, relatively
narrower, with different pronotal base and
slightly different elytral microsculpture, as
well as more lightly impressed elytral striae.
As compared with erichsoni, papua has a
wider prothorax, shallower elytral striae,
and darker prothoracic and elytral margins.
The closest relative of papua may be
ruficollis ( above ) , but the latter is bicolored
and always (in the specimens before me)
has the sutural angles subdenticulate or at
least angulate, while papua is dark, with
sutural angles usually blunted, rarely sub-
denticulate. The difference in sutural
angles is not absolute but is enough to sug-
gest that the color difference is specific.
Penfagonica esfriafa n. sp.
Description. Form of large Penta^onica;
black or piceous, legs dark, antennae in-
cluding basal segments pale; entire upper
surface with c. isodiametric microsculpture,
at most slightly transverse on elytra. Head
0.83 and 0.82 width prothorax; details of
mouthparts as in papua. Prothorax: width,
length 1.76 and 1.78; margins posteriorly
connected by a basal loop. Elytra: width
elytra prothorax 1.71 and 1.74; sutural
angles usually finely denticulate; striae ef-
faced or at most faintly indicated; dorsal
punctures, if present, minute, almost un-
detectable. Secondary sexual characters as
in preceding species (papua). Measure-
ments: length 4.9-6.2; width 2.1-2.6 mm.
Types. Holotype c^ (Bishop Mus.) and
1 9 paratype from Eliptamin Vv., N-E.
N. G., 1200-1350 m, June 19-30, 1959
(W. W. Brandt); and additional paratypes
as follows. N-E. N. G.: 1, Eliptamin Vy.,
1350-1665 m, June 23-30, 1959 {\Y. W.
Brandt, Bishop Mus.); 1, Wau, Morobe
Dist., 1200 m, Jan. 3-4, 1963 (Sedlacek);
1, "No. 14," Umi R., Markham Vy., 480 m,
Nov. 11, 1959 (L. J. Brass, A.M.N.H.).
West N. G.: 1, Hollandia, Dec. 15, 1944
(Hoogstraal, M.C.Z.); 1, Star Mts., Sibil
Vy., 1245 m, Oct. 18-Nov. 8, 1961 ( S. & L.
Quate, Bishop Mus.), taken in Malaise
trap; 3, Japen Is., Mt. Baduri, 1000 ft. (c.
300 m), Aug. 1938 (Cheesman). Some
paratypes in M.C.Z., Type No. 31,496.
Measured specimens. The 6 holotype and
1 9 paratype from Eliptamin Vy.
Notes. P. c.striata is the same size and
nearly the same form as papua but slightly
broader, with differently colored append-
ages, usually denticulate rather than blunted
sutural angles, and virtually no elytral
striae. The latter character, as well as
color, differentiat(\s estriafa from ruficollis.
PARASCOPODES n. gen.
Diagnosis. Form Scopodes-Mke. Head:
eyes enormous; labrum transverse-quadrate,
6-setose; mentum \\'ith basal suture in-
196 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
dicated, and with broad, short, more or less
emarginate tooth; hgula not swollen, 2-
setose; paraglossae much longer than ligula,
with apices narrowly rounded. Prothoia-x
weakly subpedunculate, angulate at sides,
2-setose each side. Elytra with entire, regu-
lar striae; 3rd intervals very inconspicuously
3-punctate. Secondary sexual characters: 6
front tarsi narrow, 2-seriately squamulose
below; i copulatory organs as figured
(Fig. 178); most other characters as in
Scopodes (below).
Description. See Diaiinosis (above) and
Description of the single species (below).
Type species. Scopodes cyaneus Sloane.
Generic distribution. As of the single
known species: eastern New Guinea and
North Queensland, Australia.
Notes. Although the Sro/>or/c.s-like form
and enormous eyes are striking specializa-
tions, the labrum, mentum, ligula, elytra,
and perhaps the S tarsi of Farascopodes
cyaneus seem generalized within the tribe
and suggest that the insect may be a
superficially specialized relict of a primitive
stock from which both Fentaiionica and
Scopodes may have evolved.
Farascopodes cyaneus (Sloane)
Scopodes cyancii.s Sloane 1907, Proc. Linncan Soc.
New South Wales 32, p. 380.
Description. With characters of genus;
form as in Figure 120; color above blue, more
or less purplish on elytra, below piceous;
appendages testaceous, mouthparts in part
browner; shining, reticulate microsculpture
virtually absent on head, light, irregular,
usually slightly transverse on pronotum and
elytra. Head 1.36, 1.32, 1.31 width pro-
thorax; 2 setae over each eye; Iront smooth
at middle, with a deep groo\'e each side.
Prothorax: width length 1.19, 1.16, 1.23;
margins narrow, each with 2 setae on small
projections; disc strongly convex; middle
line and trans\-erse impressions sharply im-
pressed, latter irreguhuK subpunctatc; sur-
face of disc smooth excei^t \\ illi laiiit liaiis-
verse strigulation toward sides. I'Jyhd:
width elytra prothorax 1.72, 1.72, 1.65;
striae entire, regular, coarsely punctate;
intervals regular, flat or slightly convex,
3rd with 3 very inconspicuous dorsal punc-
tures (see Notes, below). Inner icings
dimoiphic: short in 1 i from Dobodura,
full in all other specimens. Lower sui^ace
sparsely pubescent; base of abdomen on
each side ( under bases of femora ) with c.
6 parallel grooves that may form a stridu-
lating organ. Legs slender; 4th hind-tarsal
segments shallowly emarginate; 5th seg-
ments with accessory setae; claws simple.
Secondary sexual characters: i tarsi as de-
scribed under genus; 6 with 1, 9 2 setae
each side last ventral segment. Measure-
nients: length 3.7-4.0; width 1.4-1.6 mm.
Type. From Kuranda, North Queens-
land, Australia; should be in Sloane Coll.,
Canberra, but I was not able to find it
there in 1957.
Occurrence ifi New Guinea. Papua: 3,
Dobodura, Nhir.-July 1944 (Darlington); 1,
Misima Is. ("St. Aignan") (ex coll. F. C.
Morrell, British Mus.). Also 1, "New
Cuinea, Sayer" (Sharp Coll., British Mus.).
Measured .specimens. A pair ( c^ 9 ) from
Dobodma and a 9 from N. of Mareeba,
Australia (see below); figures listed in this
order.
Notes. Besides the specimens from New
Cuinea, I have 1 9 taken by myself N. of
Mareeba (about 20 miles SW. of the t>'pc
locality). North Queensland, Australia, Feb.
1958, in flooded grassland. 1 can lind no
significant difference between the New
Cuinean and Australian specimens, but oi
course I have not been able to compare 6
characters.
\u a footnote to the original description.
Sloane sa\s that he eould liiid onl\' 1
( fine ) dorsal puncture on the el\tra oi the
t\pe. However, under a good stereoscopic
inieioseope with good light, 1 lind what
appear to be .3 punctures neaiK e\('iil\
spaced along the length of each 3rd inter\al
in both New (iuinean and Australian in-
(li\ idnals.
The Carabid Beetles of Ne\\' Guinea
Darlington
197
The habitat of this species, in or under
grass at low altitudes, is different from that
of any Scopodes in New Guinea. If this
insect is a relict of an ancestral stock from
which Pcntap.onica and Sco))odcs have
evolved, as I ha\e suggested under the
genus, it may still be in an ancestral habitat
from which Pentag,onica may have invaded
low vegetation, and Scopodes, habitats on
the ground.
Genus SCOPODES Erichson
Krichson 1842, Archiv fiir Naturgeschichte 8,
Band 1, p. 123.
Sloane 1903, Proc. Linnean Soc. New South Wales
28, pp. 637-638 (key to Australian species).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1504 (see for additional references, synonymy,
and list of species ) .
DiaiJ^nosis. Form (Fig. 121) character-
istic, with eyes very large and sides of
prothorax usually angulate; mentum not
toothed; ligula swollen, club-like, longer
than paraglossae, 2-setose; i front tarsi
with soles of numerous close-packed slender
squamae not in 2 series. For additional
diagnostic characters see under the tribe
and Key to Genera of Pcntaiionicini.
Description. None required here, but
note wings full or reduced or dimorphic in
different Australian species, full in all New
Guinean species except altus, in which
reduced.
Ty})e species. Scopodes hoops Erichson,
of southern Australia.
Generic distribution. Australia, New
Zealand, New Guinea, Java; and 1 species
(not seen by me) described from New
Caledonia.
Notes. This genus divides into 2 groups.
The typical group includes all the Austra-
lian and New Zealand species, 1 species
(altus) at very high altitudes on the Snow
Mts. in New Guinea, and 1 ( irregidaris
Andrewes) on mountains in Java. The
other New Guinean members of the genus
form a distinct, endemic group of relatively
compact, often brightly colored species oc-
curring mostly at moderate altitudes in the
mountains. These species can be arranged
in what seems to be an evolutionary se-
quence involving change of form, loss of
elytral striae, simplification and reduction
of sculpture, and intensification of color,
but the lines of evolution have probably
been complex. In the last species of the
Key (adonis. Fig. 123), which is at the end
of the apparent evolutionary sequence, the
Scopodes form is much modified (the pro-
thorax being long and narrow and without
lateral angulations, and the elytra being
oval ) ; the elytral striae and sculpture have
almost disappeared, and the color is deep
purple.
These beetles live in the ground or in
logs. They are probably diurnal: most of
them are winged, but they apparently do
not come to light at night. Evelyn Ghees-
man, in The Land of the Red Bird ( London,
Herbert Joseph Ltd., no date, p. 134), says
apparently of Scopodes cheesmani (which
I describe below), "Other insects were
very tiny violet beetles, which appeared
from inside the log, out of one of its many
passage ways, and ran about on the sur-
face." This was between 3400 and 4500 ft.
(c. 1380 m) altitude in the rain-forested
Cyclops Mts.
In the following descriptions, the width
of prothorax includes the tubercles. The
frontal sulci are counted between the eyes
just behind the anterior supraocular setae.
The sculpture of the front of the head, espe-
cially of the clypeus, is variable, heavily
impressed or almost obsolete in different
individuals of single species from single
localities, and I have therefore not de-
scribed it in detail in most cases.
Key to Species of Scopodes of New Guinea
1. Prothorax with 2 setae each side, the pos-
terior on conspicuous dentiform processes;
(depressed; elytra with c. complete striation
indicated) ( p' 198) altufi
- Prothorax with only 1 seta each side, c. %
of prothoracic length from apex, without
posterior-lateral setae or processes - 2
198 Bulletin Museum of Comparative Zoology, Vol. 137. No. 1
2. Elytra with conspicuous irregular sericeous
pattern aiul extensively but irregularly
striate 3
- Elytra without conspicuous sericeous pat-
tern 4
3. Color bronze, elytral foveae conspicuously
bine, legs pale (p. 199) tafa
- Color dark bronze or greenish, eh tral foveae
not conspicuoush' contrasting, legs dark (p.
199) ..--' cJihithti
4. Color usually bronze, never primarily bright
blue or purple 5
- Color primarily blue or purple 7
5. Elytra with striae indicated at least anteri-
orly 6
- Elytra without striae (p. 201) sim))lcx
6. Extensively snbstriate (p. 200) icilsoiii
- Striae indicated only at base of elytra (p.
200) Inisdlis
7. Prothorax with lateral margins (p. 201 )
checsiiuini
- Prothorax without lateral margins (p. 201)
adoni.s
Scopodes alius n. sp.
Description. With characters of genus;
form (Fig. 121) relatively depressed, with
humeri narrowed; black, appendages dark;
rather shining, but complexly sculptured
as described below. Head 1.26 and 1.23
width prothorax; labrum long, strongly
rounded, with 4 decurved setae anteriorly,
the inner setae much shorter than the outer
(setae sometimes broken off); front with
c. 20 or more fine sulci, latter subparallel
at sides and base but connected and partU
transverse at middle ol head; front anteri-
orly and clypeus and labrum more finel\
and closely longitudinally sculptured in
S 6 , sculpturing reduced in 9 (see Notes).
Prothora.x: width length 1.24 and 1.27;
sides margined, each with 2 setae, at an-
gulation c. ':'. from apex and on conspicuous
triangular process b(4()re base; disc with
weak median longitudinal imiiression and
strong subbasal and subapical transverse
imprt\ssions; surface of disc covered vvith
rather line, complex, more or less anas-
tomosing sculptuic. Elytra: width elytra/'
prothorax 1.69 and — (elytra too spread to
measure in 9 ); striae indicated ])\' imdula-
tions of the surlace bul not sliarpK dclincd.
slightly irregular; intervals slightly convex,
3rd with 3 moderate foveae; elytral surface
with rather light irregular reticulate micro-
sculpture. Inner tt/ng.s- reduced to c. Vs
length of elytra. Measurements: length c.
3.3-3.5; width c. 1.3 mm.
Types. Holotype S (Leiden Mus. ) from
Scree Vy. Camp, Snow Mts., West N. G.,
3600 m, Sept. 19, 1938; 1 i paratype
(M.C.Z., Type No. 31.497) from same
locality, 3800 m, Sept. 1938; 1 9 paratype
(Leiden Mus.), from Lake Habbema, Snow
Mts., 3250-3300 m, late July-Aug. 1938 (all
specimens collected by Toxopeus ) .
Measured specimens. The 6 holotype and
9 paratype.
Notes. This species differs from all other
Scopodes known from New Guinea in its
depressed form and in possessing 2 pairs
of lateral prothoracic setae. In these ways it
resembles some Australian species including
the type of the genus, Scopodes hoops
Erichson, but (dttis is more shining than
Jjoops, with better defined pronotal sculp-
ture, and other minor differences of detail.
S. altus resembles also irre^idaris Andrewes
of Java but is black rather than aeneous,
with elytra less narrowed anteriorly and
more distinctly striate, as shown b\ com-
parison with a specimen ol irreu^ularis from
"G. Tengger, Java, l^rescher." Of the 3
specic^s just discussed, the Australian hoops
has dimorphic wings; the New Guinean
and Javan specie's, reduced ones; but they
all are probabK derived Irom a winged
ancestor that dispersed i)\ I light. The
distribution of this group ol Scopodes in
the Malax' Archipelago is comparable to
that of Meeyclothorax. which also occurs
primariK' in Australia but is represented at
verv high altitudes on the Snow Mts. ol
New Guinea and on the mountains ol Java
(Darlington, Hull. M.C.Z. 126. 1962, pp.
505-507). S(H' also Mieroferouia, [vige 18
ol my i^rescut pajier.
Whether the dillcrence in sculpture of
clypeus and labrum of the 6 S (Scree Vy.)
and the V (Lake llabbema) is sexual.
The Carabid Beetles ok New Guinea • DarUnglon
199
geographic, or individual cannot be decided
without more material.
Scopodes fafo n. sp.
Description. With characters of genus;
form compact, convex; aeneous, labrum
dark, dorsal and lateral foveae of elytra
blue, appendages pale, antennae browner
distally; head and pronotum shining, sculp-
tured as described (below), elytra irregu-
larlv sericeous with irregular reticulate
microsculpture. Head 1.19 and 1.20 width
prothorax; labrum rounded, 6-setose; front
with c. 8 longitudinal sulci abbreviated an-
teriorly and running into coarse rugosity
posteriorly; labrum more finely longitudi-
nally rugose. ProtJwrax: width length
1.26 and 1.27; sides margined, each with
seta-bearing puncture on triangular process
at angulation c. % from apex; disc with
coarse rugosity transverse posteriorly but
more confused anteriorly. Elytra subquad-
rate; width elytra prothorax 1.83 and 1.77;
several striae indicated on disc but striation
obsolete externally and apically; 3rd in-
terval conspicuously 3-foveate. Measure-
ments: length 3.5-4.0; width 1.4-1.7 mm.
Ti/pes. Holotype 9 (British Mus.) and
4 paratypes (2 in M.C.Z., Type No. 31,498)
all from Mt. Tafa, Papua, 8500 ft. (c.
2600 m), Feb. 1934 (Cheesman).
Additional material. One $ , Wau, Bull-
dog Rd., N-E. N. G., 2400 m, May 31, 1962
( Sedlaceks ) .
Measured specimens. The c^ holotype and
1 9 paratype.
Notes. See preceding Key for characters
distinguishing this from related species.
The specimen from Wau is slightly larger
than the types, with narrower humeri (but
still with large folded inner wings ) and less
distinct elytral striation. Additional ma-
terial may show it to be a distinguishable
form.
Scopodes chimbu n. sp.
Description. With characters of genus;
form compact; color dark, subaeneous or
greenish, elytral foveae not contrasting,
appendages dark except antennae paler
basally; head and pronotum sculptured but
with irregular reticulate microsculpture.
Head 1.15 and 1.22 width prothorax; labrum
rounded, 6-setose; front with c. 7 longitu-
dinal sometimes slightly sinuous sulci
shining, elytra irregularly sericeous and
running into coarse rugosity posteriorly and
sometimes anteriorly. Prothorax: width/
length 1.25 and 1.21; sides margined, each
with 1 seta-bearing puncture, on tubercle
at angulation c. % from apex; disc coarsely
rugose, the rugosity in general transverse
but somewhat confused especially anteri-
orly. Elytra subquadrate; width elytra/
prothorax — and 1.83 ( elytra of c5 too spread
to measure); several striae indicated on
disc, outer striae fainter or obsolete; 3 con-
spicuous foveae on each 3rd interval. Mea-
surements: length 3.5-4.4; width 1.4-1.7
mm.
Types. Holotype $ (M.C.Z., Type No.
31,499) and 2 (99) paratypes from
Chimbu Vy., Bismarck Rge., N-E. N. G.,
5000-7000 ft. (c. 1500-2135 m), Oct. 1944
( Darlington ) .
Additional material. N-E. N. G.: 2, Mt.
Mis(s)im, Morobe Dist. (1 specimen at
6400 ft. = 1950 m) (Stevens, M.C.Z.); 1,
Joangang, 500 m, Apr. 7-8, and 1, Tumnang,
1400-1600 m, Apr. 14-15, both on Mongi
Watershed, Huon Pen. ( 1955, E. O. Wilson,
M.C.Z.); 1, Saruwaged (Salawaket) Rge.,
upper Bunbok Vy.. 1800-2000 m. May 1955
(E. O. Wilson, M.C.Z.); 1, Sepalakembang,
Salawaket Rge., 1920 m, Sept. 12, 1956 (E.
J. Ford, Jr., Bishop Mus.); 1, Kepilam,
2420-2540 m, June 21, 1963 (Sedlacek); 1,
Finisterre Rge., Saidor, Matoko, Aug. 29-
Sept. 5, 1958 (W. W. Brandt, Bishop Mus.).
Measured specimens. The 6 holotype and
1 9 paratype.
Notes. This species varies somewhat from
locality to locality, but I do not have enough
material to distinguish geographic forms.
Except that my specimens were taken on
the ground in the more open part of the
200
Bulletin Miiscinu of Comparative Zoology, Vol. 137, No. 1
Chimbu Valley, I can say nothing about
their habits.
Scopodes wilsoni n. sp.
Description. With characters of genus;
compact, convex; color dark, subaeneous,
elytral foveae not contrasting, appendages
dark, antennae paler basally; shining, head
and pronotum coarsely sculptured, elytra
irregularly microreticulate but without con-
spicuous sericeous pattern. Head 1.25 and
1.24 width prothorax; labrum narro\\'ly
rounded, 6-setose; front with c. 8 longitu-
dinal sulci running into coarse rugosity pos-
teriorly and usually anteriorly. Frothorax:
width/length 1.23 and 1.18; sides margined,
each with seta-bearing puncture on tuber-
cle at angulation c. % from apex; disc
coarsely rugose, the rugosity transverse but
somewhat irregular and varying in depth,
and disc also variably punctulate. Elytra
quadrate; width elytra/prothorax 1.91 and —
(elytra too spread to measure in second
specimen); all or several striae indicated
for much of length; 3 conspicuous foveae on
or near each 3rd interval. Measurements:
length 3.4-4.1; width 1.4-1.7 mm.
Types. Holotype $ (M.C.Z., Type No.
31,500) from Nganduo to Yunzain, Mongi
Watershed, Huon Pen., N-E. N. G., 1000-
1500 m, Apr. 6, 1955 (E. O. Wilson); and
following paratvpes (all i 6 ) from N-E.
N. C: 1, Nad/ab, Markham Vy., July 13,
1944 (K. V. Krombein, U.S.N. M.), this
specimen further labeled "E. fork Ngafir
Cr. 100()-3()00 ft. nativc> trail"; 1, Mt.
Mis(s)im, Morobe l^i.st., 6400 Ft. (1950 m)
(Stevens, M.C.Z.); 1, Wau, 1300 m, Dec.
10, and 1, same locality, Nami Ck., 1750 m,
Aug. 12 (both 1961, Sedlaceks).
Addilional nuiterial. Fapua: 1, Owen
Stanley Rge., Goilala, Bome, 1950 m, Apr.
30-Mav 2, 1958 (W. \V. Brandt, Bishop
Mus.).' West IS. (;.: J, Battau Camp, 1150
m, Feb.-Mar., and 1, Sigi (lamp, 1500 in,
Feb. 26, both 1959 (Neth. Ind. -American
[Snow Mts.] Exp.; Toxopeus).
Measured specimens. The A holotype and
9 paratype from Nad/ab.
Notes. As in the case of chimbu (pre-
ceding), this species appears to vary geo-
graphically, but my material is too limited
to justify describing geographic forms.
Scopodes basalis n. sp.
Description. With characters of genus;
very compact, convex; dark green or aene-
ous, elytral foveae not contrasting, labrum
and legs dark, antennae yellowish brown;
shining, reticulate microsculpture faint or
absent e\en on elytra. Head 1.22 and 1.20
width prothorax; labrum narrowly rounded.
6-setose; front with c. 7 longitudinal sulci
sometimes abbreviated anteriorly and pos-
teriorly. Trothorax: width length 1.21 and
1.25; sides margined, each with seta-bearing
puncture on tubercle at angulation c. V:i
from apex; disc coarsely transversely rugose
(rugosity sometimes only lightK" impressed)
and rather sparsely fineh' punctulate.
Elytra quadrate; width elytra/prothorax
1.79 and 1.82; striae absent or \irtuall\- so
except deeph' but variably impressed at
base, sometimes only on basal declivit\-; 3
conspicuous punctiform foveae on positions
of 3rd interxals. Measurements: length c.
3.3; width c. 1.7 mm.
Types. Holotype A (M.C.Z., Type No.
31,501) from Joangang, Mongi Watershed,
Huon Pen., N-E. N. G., 500 m, Apr. 7-8,
1955 (E. (). \\'ilson); and following para-
types from N-E. N. (,.: 1, Saruwageil
(Salawaket) Rge., upper Bunbok \'v., 2300-
3200 m. May 29-31 (E. (). Wilson, M.C.Z.),
"mossy forest"; 1, Mt. Mis(s)im, Morobe
Dist. (Stevens, M.C.Z.); 1, TorricelH Mts..
Siaute, sea level, Nov. 9-17, 1958 (\\. W.
Brandt, Bishop Mus. ).
Measured sjx'cimens. The s holot\ jie and
9 paratyjie Irom Saruwaged Bge.
Notes. S. basalis diHers from iriisinii not
oiiK in reduction ol cKtral striation bnt
also in more pnnelilorm cKtral loxcae and
\irtnal absence ol icticnlatc inicrosculi')turc>
of eKtia. 'Ilic oeiui rcnce ol an indixidnal
near sea IcncI is uiinsnal in this genns in
New (iuinea.
The Carabid Beetles of New Guinea • Darlington
201
Scopodes simplex n. sp.
Description. With characters of genus;
compact, convex; aeneous black, elytra
sometimes faintly purplish, front anteriorly,
clypeus, and labrum ])right aeneous, ap-
pendages dark, base of antennae paler;
shining, elytra lightly irregularly microreticu-
late. Head 1.25 and 1.25 width prothorax;
front with 7 sulci running into coarse
rugosity posteriorly and sometimes anteri-
orly. Prothorax: width/length 1.19 and
1.22; sides margined, each with seta on
tubercle c. Mi from apex; disc lightly trans-
versely wrinkled, ± punctulate ( variable ) .
Elytra subquadrate; width elytra prothorax
1.90 and 1.92; disc virtually estriate, with 3
conspicuous punctiform foveae near posi-
tion of each 3rd interval. Measurements:
length 3.8-4.2; width 1.5-1.7 mm.
Types. Holotype c^ (M.C.Z., Type No.
31,502) and 2 paratypes from Nganduo to
Yunzain, Mongi Watershed, Huon Pen.,
N-E. N. G., 1000-1500 m, Apr. 6, 1955 ( E.
O. Wilson); and additional paratypes as
follows. N-E. N. G.: 1, Gemeheng, Mongi
\\'atershed, Huon Pen., 1300 m, Apr. 11-13,
1955 (E. O. Wilson, M.C.Z.); 1, vie. Xad-
zab, July 1944 (Darlington); 2, Sattelberg
(British Mus., ex Coll. Hauser). West
N. G.: 8, Wissel Lakes, Moanemani, Kamo
Vy., 1500 m, Aug. 14, 1962 (Sedlacek); 2,
Wissel Lakes, Enarotadi, 1500 m, Aug. 14,
1962 (Sedlacek).
Measured specimens. The i holotype and
1 9 paratype from Nganduo to Yunzain.
Notes. This species, like the preceding
( hasalis ) , occurs at relatively low altitudes,
sometimes down almost to sea level (at
Nadzab ) .
Scopodes cheesmoni n. sp.
Description. With characters of genus;
form (Fig. 122) compact, convex; blue-
purple (in general blue with elytra puiple
with foveae bluer); shining, reticulate
microsculpture faint or obsolete even on
elytra. Head 1.31 and 1.35 width prothorax;
front with 7 sulci running into coarsely
rugose areas posteriorly and usually anteri-
orly. Frothorax narrower and less angulate
than usual; width/length 1.13 and 1.18;
sides margined, each with seta on small
projection c. M', from apex; disc coarsely
transversely sulcate or wrinkled, ± punctu-
late especially anteriorly. Elytra subquad-
rate, slightly narrowed toward base; width
elytra/prothorax 1.98 and 2.09; disc estriate,
with 3 conspicuous punctiform foveae on
position of each 3rd interval. Measure-
ments: length 3.9-4.4; width 1.6-1.8 mm.
Types. Holotype $ (British Mus.) and
2 paratypes from Mt. Lina, Cyclops Mts.,
West N. G., 3500 ft. (1067 m). Mar. 1936
( Cheesman ) ; and additional paratypes as
follows. West N. G.: 4, Cyclops Mts.,
3400-4500 ft. (c. 1040-1370 m). Mar. 1936
(Cheesman); 3, Rattan Camp, 1150 m,
Feb.-Mar., and 1, Sigi Camp, 1350 m. Snow
Mts., Feb. 28, 1939 (Toxopeus); 1, Bivak
36, 1220 m, July 30, and 1, Bivak 39, 1300
m, June 30, 1959, Star Rge. (Neth. New
Guinea Exp., Leiden Mus.). N-E. N. G.:
1, Chimbu Vv., Bismarck Rge., 5000-7500
ft. (c. 1500-2300 m), Oct. 1944 (Darling-
ton); 1, Eliptamin Vy., 1665-2530 m, June
23-30, 1959 (W. W. Brandt, Bishop Mus.).
Some paratypes in M.C.Z., Type No. 31,503.
Measured specimens. The i holotype and
1 9 paratype from Mt. Lina.
Notes. Although not very different in
form and basic structure from the several
preceding compact species, cheesmani does
suggest transition toward the following
(adonis).
The habits of this species are suggested in
Notes under the genus.
Scopodes adonis n. sp.
Description. With characters of genus;
form ( Fig. 123 ) less compact, with longer
prothorax and more oval elytra, than in
other members of the "New Guinean group"
of Sco})odes; purple; front anteriorly,
clypeus, and labrum cupreous; legs dark;
antennae with basal and outer segments
brown, segments 2-5 paler; shining, upper
202 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
surface without reticulate microsculpture,
sparsely punctulate. Head 1.37 and 1.32
width prothorax; front with c. 7 slightly
unequal sulci behind level of anterior
supraocular punctures, the sulci abbreviated
anteriorly and cur\'ing toward sides pos-
teriorly; neck irregularly rugose. Prothorax
oval long; width length 0.93 and 0.96;
sides not margined, not angulate, each with
seta-bearing puncture c. Va from apex; disc
strongly convex, with fine but distinct mid-
dle line, weakly transversely wrinkled.
Elytra suboval; width elytra prothorax 2.13
and 2.17; humeri narrower and more
broadly rounded than in other species of
group; outer-apical angles obtusely angu-
late, sutural angles acute; striae absent; 3
minute, inconspicuous seta-bearing punc-
tures on position of each 3rd interval. Mea-
siirement.s: length 4.7-4.9; width 1.7-1.9
mm.
T\j])es. Ilolotype i (Bishop Mus.) and
8 paratypes (some in M.C.Z., Type No.
31,504) from Torricelli Mts., Mokai Village,
N-E. N. G., 750 m, Dec. 8, 16 (holotype,
Dec. 8), 1958 (W. W. Brandt); and 2 para-
types, Torricelli Mts., Mobitei, 750 m, Feb.
28-Mar. 4 and Apr. 1-15, 1959 (W. W.
Brandt, Bishop Mus.).
Measured speeimcn.s. The i holotype and
1 9 paratype from Mokai Village.
Notes. This very distinct species is dis-
tinguished from all others known to me by
form, prothorax long with margins obsolete,
elytra oval with acute sutural angles, elytral
foveae reduced, and color. However, as
noted in discussion under the genus, it prob-
ably does not represent a separate stock in
New Guinea but seems to be at the end of
an evolutionary sequence which includes
other New Guinean species.
Tribe HEXAGONIINI
Csiki 19.'32, Colcop. Cat., (;aial)itl;K', Ilaipalinac 7,
p. 1506 (sec tor s>n()n\niy and additional rclcr-
enccs).
flcxofioniitue Jcanncl 1948, (lolrop. C'aiahiqiics dc
la i^ogion Malgadic, I'ait 2, p. 7.59.
This is a small tribe (4 genera) of char-
acteristically formed ( Fig. 124 ) , subparallel,
often flattened carabids at least some of
which are specialized to live under the
leaf sheaths of plants. In this tribe the
inner lobe of the maxilla has a slender,
Diocahle apical segment that is diagnostic,
occurring in no other Carabidae except the
Cicindelinae.
The tribe is confined to the Old-World
tropics. The genus Hexa^onia (below) is
widely distributed there; 1 additional genus
occurs in the Oriental Region; 2 more, in
Madagascar.
Genus HEXAGON/A Kirby
Kirby 1825, Trans. Linnean Soc. London 14, p.
563.
Csiki 1932, Coleop. Cat., Cavabidat', Ilaipalinae 7,
p. 1506 (see for synonymy and additional refer-
ences ) .
Jeannel 1948, Coleop. Carahiques de la Region
Malgache, Part 2, p. 759.
Basilewsky 1948, Bull. Mus. Hist. \at. Belgian
Congo 24, p. 3 (African species).
Diaiinosis. See under tribe, of which this
is the only genus represented in New
Guinea.
Deseri))t}on. None required here.
Ty))e s))eeies. II. terniiiuifa Kirby, of SE.
Asia.
Generie distribution. Tropical .\sia and
islands to New Guinea and northeastern
Australia; Africa, Madagascar.
Notes. Members of this genus are rather
diverse in the Oriental Region including
the Philii)pines, but ()nl\ I spc^cies group
extends to New (Guinea and Australia.
ProbabK' bee;uise the\- oceup) an unusual
niche and perhaps also beeausi' the\' are
diurnal and nia\ not ll\ to light, these
insects are rarel\ eolleeled. The 1 New
Guinean and the 1 ( nnd(\scrib(Kl ) .Austra-
lian s|ieeies are eaeli known Ironi a single
collection made b\ nusell, by breaking
down tall grass and other a([uatic \-egeta-
tion into water and picking np the beetles
;is the\ came to the surhice.
The Carabid Beetles of New Guinea • Darlington 203
Hexagonia popua n. sp.
Description. With characters of tribe;
form as in Figure 124; head black; pro-
notum piceous, reddish at base and apex;
elytra red in anterior 1-2 or more, black
posteriorly, the black area extending farther
forward on inner than on outer part of
elytra; lower surface irregularly reddish and
piceous; appendages brown; shining, reticu-
late microsculpture faint or absent on head
and pronotum, imperfect on elytra; head
and pronotum irregularly punctate. Head
1.05 and 1.05 width prothorax; front semi-
circularly impressed each side; neck deeply
transversely impressed. Prothorax subcor-
date; width length 1.06 and 1.09; base/apex
not calculated (because prothorax rounded
into neck without distinct anterior angles);
margins narrow, each with seta c. % from
apex, without posterior seta; disc with mid-
dle groove deeply impressed and subpunc-
tate, other impressions irregular and weak.
Elytra: width elytra/prothorax 1.66 and
1.67; striae well impressed, punctate; inter-
vals slightly convex, 3rd with 3 or 4 con-
spicuous seta-bearing punctures, 5th with 1
such puncture c. ^.i from apex on outer edge.
Lower surface: head below transversely
rugulose; prothorax below subrugosely
punctate. Inner wings full. Legs moderate;
tarsi wide; 4th tarsal segments very deeply
emarginate, long-lobed; claws simple, not
toothed. Secondary sexual characters: S
tarsi not or not much modified, without
special squamules; i with 2, 9 3 short
setae near apex each side last ventral seg-
ment. Measurements: length 7.2-7.8; width
2.2-2.4 mm.
Types. Holotype S (M.C.Z., Type No.
31,505) and 12 paratypes all from Aitape,
N-E. N. G., Aug. 1944 (Darhngton).
Measured specimens. The c^ holotype and
1 ? paratype (sexes of these specimens
determined by dissection).
Notes. This new Hexagonia is probably
related to lucasseni van der Poll of Java but
is slightly more slender and much darker,
with pronotum piceous (red in lucasseni)
and elytra more extensively black posteri-
orly. One or more similar but apparently
undescribed species occur in the Philip-
pines.
The habitat of the species is noted under
the genus.
Tribe ODACANTHINI
Sloane 1917, Proc. Linnean Soc. New South Wales
42, p. 413.
1923, Proe. Linnean Soe. New South
Wales 50, p. 30.
Jedlieka 1963, Ent. Abhandlungen 28, p. 488.
Habu 1967, Fauna Japonica, Carabidae, Tnui-
eatipennes Group, p. 13.
Odacantliidae Jeannel 1948, Coleop. Carabiques
de la Region Malgaehe, Part 2, p. 745.
Odacanthinac Basilewsky 1953, Exploration Pare
National I'Upemba, Fase. 10, Carabidae, p. 108.
CoIUuriui Csiki 1932, Coleop. Cat., Carabidae,
Ilaipalinae 7, p. 1517 (see for .synonymy and
additional references ) .
Liebke 1938, Festschrift Einbrik Strand 4, pp. 37-
141.
Most members of this tribe have a char-
acteristic form (Figs. 125-131), with pro-
thorax very long and narrow, usually much
narrower than head. Technical characters
of the tribe are given by authors cited
above. The tribe is well represented in
the tropics of both hemispheres, less well
represented in most temperate regions, but
several remarkable endemic genera occur
in Australia.
Liebke ( 1938 ) has published a useful
generic classification of this tribe for the
world. His classification is, however, artifi-
cial, as shown by the failure of some of my
new New Guinean species to fit into it.
The form of the 4th hind-tarsal segments,
used by Liebke in the first couplet of his
key, is a particularly unsatisfactory generic
character. Form of the 4th hind-tarsal seg-
ments does characterize some genera, but
it is extremely variable in others. See, for
example. Notes under Dicraspeda in the
following pages. However, I cannot under-
take to revise the generic classification in
dealing with the few members of the tribe
that occur in New Guinea, except in the
case of primarily New Guinean genera.
204 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Ecologically, Odacaiithini are active,
winged carabids some of which live in
foliage and some on the ground. Of the
New Guinean forms, Dicraspeda ( includ-
ing Philcmonia and Macrocentra) lives in
understory foliage of rain forest; Clarcncia
and some CoUiuris, in or under low vegeta-
tion or dead leaves in wet places; Casnoideo,
among reeds and in other vegetation in
water; Eudalia (in Australia), Dohodura,
and I think also Lachnothorax, in gravel
and among stones bv running water; and
Basisticu.s ( in Australia ) , on the ground
in relatively dry places including open
Eucahjptu.s woodland.
Eight genera, 19 species of Odacanthini,
are known from New Guinea. Two or 3
stocks of CoIUuris, 1 species of Casnoidca,
and Lachnothorax have probably reached
New Guinea rather rec(uitly from the
Oriental Region. A second species of
Casnoidea and also Basi.sticus micans (which
may be related to Coditiris) are shared
with Australia. Cdarcncia and Eudalia are
shared with Australia and may be of Aus-
tralian origin. Monotypic Dohodura is con-
fined to New Guinea but may be derived
from Eudalia. And Dicra.s))eda {.sensu lato)
has radiated chiefly in the rain forests of
New Guinea, where 6 diverse species now
exist.
Key to Genera of Odac:antiiini of
New Guinea
1. Lateral niaiyins of protlinrax iDtomplcte
(i)i(l 4tli liiiul-tarsal sciiiiK'iils \ci>' lonij;-
lobc'd; strikinjfly bicokiied, bkick (or jiiccous)
and red ( p. 207 ) (.Uisnoidca
— Not as above in one oi' more ways; nsually
not blaek-and-red l)ieoIore(l (olisenrely so
in lidsislicns ) 2
2. Antennae with 3rd se,unients \er_\ h)njf, c.
% longer than 4th segments (p. 209) _..
Cldicncid
— Antennae with ■ivd segments shorter, ('{jual
to or not more tlian 'A longer tlian 4t]i
segments 3
3. Head with fine eosta eaeh side above eye
ami pronotum elianneled at sides and mid-
(Ue; ( l)ase ot elytra not i(un\clti pnnetate-
striate) (yi. 210) l^icraspcdd
— Not as above in one oi more ways 4
4. Body pnbeseent ([i. 214) I .(kIiiioIIidiiix
- Not pubescent 5
5. Eyes not margined on inner edges by costae
and front smooth 6
- Eyes usually margined by costae, or if
(rarely) not margined, front so coarsely
pimetate that supraocular costae are indis-
tinct 7
6. Elytra not spined (p. 214) EiidaUd
- Elytra spined (see also Notes under this
genus) (p. 215) ... Dohodura
7. Side margins of prothorax absent or incom-
plete or ( if margins nearly complete ) pro-
notum not channeled at middle; elytra not
\erv differentlv sculptured at base and apex
(p.' 205) _._! Colliuris
- Side margins of prothorax complete and
pronotum channeled at middle; elytra
coarsely punctate-striate in anterior %,
smooth posteriori)' ( p. 20(S ) Basisticiis
Genus COLLIURIS Degeer
Degeer 1774, Mem. Hist. Insectes 4, p. 79.
Gsiki 19.32, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1.518 (see for synonymy and additional refer-
ences ) .
Liebke 1938, Eestschrift Endirik Strand 4, p. 45.
ledlicka 1963, Ent. Abhandlungen 28, pp. 489.
490.
Diuiinosis. See preceding Keij to Genera.
Deseription. None required here.
Tijpe speeies. Attelahus surinameusis
Linnaeus, of South America.
Generic distribtition. All wami regions
of the worlfl. In the Asiatic- Australian area,
species are moderately numerous and di-
verse in southern Asia and the western
Malay Archipelago, but only 4 species
( representing 3 indc^pcMident stocks ) have
been foiuid in New (Guinea. According to
Liebke, the genus does not reach Australia,
but I found 3 species of it ( representing 2
ol the same stocks that are in New Guinea)
in North Oueensland in I95(S.
Notes. I cannot fit the New Guinean
species into T-iebkes subg(Mi(Ma satisfac-
torily. ('. ro.s.si (below) and also the Aus-
tralian C. ohscura Gastelnau ha\e \irtualK
complete lateral prothoracic margins, which
rules these species out of Colliuris entirely,
according to Li(>bke (who in fact incor-
rectly put ohscura in Dicraspeda). Never-
theless, in loiiu and other characters these
The Carabiu Beetles of New Guinea • Darlington
205
species seem to me to be closely related to
Colliuris subgenus EticoUiuris of Africa and
the Oriental Region.
Key to Species of Colliuris of New Guinea
1. Prothorax with lateral margins; intervals 3,
5, 7 with seta-bearing punctures; (pronotum
and elytral striae coarsely punctate, head
smooth) (p. 205) rossi
- Prothorax without lateral margins, or margins
reduced to sutures; only 3rd intervals with
seta-bearing punctures 2
2. Not maculate; subcylindric, elytra not more
than 2x width prothorax; entire upper sur-
face coarsely punctate (p. 205) fii.svipcnnis
- Quadrimaculate; broader, elytra more than
2X width prothorax; upper surface c. im-
punctate except for finely punctate elytral
striae 3
3. Less slender, prothoracic width/length 0.60
and 0.63; elytra not or not much impressed
(p. 206) papiiu
- More slender, prothoracic width/length 0.46
and 0.46; elytra transversely impressed near
base (see also Description) (p. 206) par
Colliuris rossi n. sp.
Description. Form (Fig. 125) of Colliuris
of Oriental fiiscipennis group; black, legs
brown with dark knees, antennae brown,
paler at base; shining, without reticulate
microsculpture. Head 1.13 width prothorax;
right mandible with acute tooth on inner
edge; a fine costa over each eye separated
from eye by groove; anterior and posterior
supraocular seta-bearing punctures present
but no other setae posteriorly; front convex,
slightly impressed at middle and anteri-
orly, impunctate; mentum with acute tooth;
ligula broad, 2-setose; palpi slender, acumi-
nate, not pubescent. Prothorax long-oval;
width/length 0.71; base apex 1.34; side
margins entire except confused by puncta-
tion at extreme base, narrow, each with 1
seta before middle; middle line light; disc
strongly convex, irregularly coarsely punc-
tate. Elytra subparallel; width elytra/pro-
thorax 2.00; apices obliquely sinuate-trun-
cate with outer and sutural angles blunt;
striae coarsely punctate, the punctures be-
coming finer posteriorly; intervals 3, 5, 7
each with 4 to 6 well spaced seta-bearing
punctures. Inner wings full. Lower surface
not pubescent, coarsely punctate anteriorly
including sides of metasternum, smooth
posteriorly. Legs normal; tarsi not pubes-
cent and not sulcate above; 4th hind-tarsal
segments shallowly emarginate. Secondary
sexual characters: 6 front tarsi narrow, 3
segments narrowly 2-seriately squamulose;
c5 last ventral segment c. semicircularly
emarginate at apex, with 1 seta each side; $
copulatory organs as in Figure 179; 9 un-
known. Measurements: length c. 6.5; width
2.0 mm.
Type. Holotype £ (Gal. Acad.) from
Finschhafen, N-E. N. G., May 7, 1944 (E.
S. Ross); the type is unique.
Notes. This new species seems closely
allied only to C. obscura (Castelnau) of
NE. Australia, but ohscura has only the 3rd
elytral intervals with seta-bearing punc-
tures. Otherwise the two species agree in
most characters including presence of
virtually entire prothoracic margins, distri-
bution of coarse punctation (head impunc-
tate, pronotum contrastingly coarsely punc-
tate), i secondary sexual characters, and
presence of an acute tooth on the right
mandible in some individuals. However,
presence of the mandibular tooth does seem
to be an individual character in ohscura:
in my series from Cairns, the tooth is well
developed in some and almost absent in
other specimens.
The generic assignment of ohscura and
rossi is doubtful. Sloane (1923, Froc.
Linnean Soc. New South Wales 48, p. 31 )
thought ohscura might go in Arame, which
is doubtful, and Liebke (1938, Festschrift
Embrik Strand 4, p. 89) put it in Dicraspeda,
which is certainly wrong. Only the pres-
ence of nearly entire prothoracic margins
prevents placing both ohscura and rossi in
Colliuris in Liebke's classification, and I
doubt if the prothoracic margins are of
generic value in this case.
Colliuris fuscipennis (Chaudoir)
Chaudoir 1850, Bull. Soc. Nat. Moscow 23, Part 1,
p. 26 {Casnoiiid) .
206
Bulletin Miiscuni of Comparative Zoology, Vol. 137, No. 1
Andrewes 1927, Ann. Mag. Nat. Hist. (9) 19, p.
106 (Odacantha).
Csiki 1932, Colfop. Cat., Carabidae, Harpalinae 7,
p. 1527 (see for synonymy, '■varieties," and addi-
tional references).
Lie1)ke 1938, Festschrift Enilirik Strand 4, p. 65,
fi,y. 27.
Tedlicka 1963, Ent. Abhandkingen 28, p. 494, figs.
190-193.
Description. None required here. Note
form rather slender; eolor bhiek with apex
of elytra reddish, legs testaceous; upper
surface including head coarsely punctate;
length c. 5% mm.
Type. From China ("Chine, Tchusan?");
in Oberthiir Coll., Paris Mus. (not seen).
Occurrence in Ncic CAiinea. Papua:
1, Lake Daviumbu, Fly R., Aug. 19-30,
1936 (Archbold Exp., A.M.N.H.).
Notes. Fiiscipennis is the oldest name
for a very common species (n- group of
closely related species previously known
from SE. Asia to Celebes and the Philip-
pines. Andrewes (1927) says of it that
fuscijx'nnis Chaudoir, punciata Nietner,
haeniorrhoidalis Motscliulsky, and jlavi-
cauda Bates "appear to dilfer very little
from each other; they may all prove to
belong to the same species, but at present
I have not the means of dc-ciding this."
(isikis, Liebke's, and Jedlicka's treatment
of some of the doubtful forms as subspecies
or varieties is not acceptable. Under these
circumstances I can only refer the New
Cuinean individual to fuscijx'nuis scnsn
l(il(K pending revision ol all related tornis.
Colliuris papua n. sp.
Description. With character's of genus;
lonn { I'ig. 126) ol ('olliuri.s, with mod-
erately broad elytra scarcely impressed be-
lore middle; black, each elytron with 2 c.
round yellow spots, centered on tth and 5th
intervals, belore middle and before apex;
apjiendages brown, antennae paler al bas(.\
moderatcK shining, reticulate microsculp-
ture hunt and r. isodiametric on front, trans-
verse on head posteriori)' and on pronofum,
indistinct on elytra. Head 1.44 and 1.11
width prothorax; 2 setae ()\cr each e\c bnt
no other setae posteriorly; front scarcely
impressed, impunctate. Prothorax long,
swollen at sides behind middle, strongly
narrowed anteriorly; width length 0.60 and
0.63; base apex 1.54 and 1.56; lateral mar-
gins reduced to sutures, each with 1 seta
before middle; disc strongly convex, base
scarcely impressed, middle line fine, surface
faintly transversely strigulose, punctate
across base. Elytra: width elytra prothorax
2.24 and 2.22; apices obliquely truncate
( slightly emarginate ) with outer angles
rounded, sutural angles scarcely blunted;
striae formed In rows of small punctures
which become minute posteriorly; 3rd in-
tervals with c. 4 seta-bearing punctures, in-
tervals 5 and 7 without punctures. Inner
winfis full. Lower surface punctate only
around front coxae and at front of meso-
sternum. Lci^s normal; tarsi not pubescent
and not sulcate above; 4th hind-tarsal s(>g-
ments shallowly emarginate. Secondary
sexual characters: A front tarsi narrow,
narrowly 2-seriately squamulose below;
last ventral segment slightK' emarginate at
apex in S , variably impressed in 9 , with
1 seta each side in c5 , 2 in 9 . Measure-
ments: length e. 5.5-6.0; width 1.6-1.8 mm.
Types. Holotype i (M.C.Z., Type No.
31,506 ) and 35 paratypes all from Dobo-
dura, Papua, Mar.-July 1944 ( Darlington ) .
Measured s])eeiniens. The ' holot\pe and
1 9 paratype.
Notes. See Notes imder the following
species, par.
Colliuris par n. sp.
Description. Similar in forTU and most
characters to preceding t papua ) but more
slender, with elytra impressed before mid-
dle; anterior eUtral spots longer antl almost
eonlined to 5th inlei\als, legs pale at base;
reticulate microsculptnic not distinct on
pronotuni. Head 1.65 and 1. 58 width \nn-
thorax. Prothorax: width length 0.46 and
0.46; base apex 1.55 and 1.56; disc trans-
\('rsel\ iinprc^ssed and constricted before
base. Elytra: width ebtra prothorax 2.60
and 2.62. ]A)irer surface as in pa))ua excei")t
The Carabid Beetles of New Guinea • Darlington 207
with row of coarse punctures each side
prosternum before coxae. Secondary sexual
characters: as for papiia. Measurements:
length c. 6.0-7.0; width 1.7-2.0 mm.
Types. Holotype $ (M.C.Z., Type No.
31,507 ) and 5 paratypes all from Hollandia,
West N. G., July-Sept. 1944 ( Darlington ) .
Additional material. N-E. N. G.: 7,
Aitape, Aug. 1944 (Darlington).
Measured specimens. The $ holotype and
1 9 paratype.
Notes. C. papua and par seem to be
allopatric representatives of one ancestral
stock, but they differ too much to be con-
sidered subspecies. C. par is the more
widely distributed: I found it at Cape
Gloucester, New Britain, in 1944, and
(1 9 ) at Lockerbie, near the tip of Cape
York, Queensland, Australia, in 1958. A
second 9 from Lockerbie is superficially
similar but much more compact and differs
in other details. I think it probably rep-
resents a 3rd, distinct species of this group
of Colliuris.
Genus CASNOIDEA Castelnau
Castelnau 1834, Etudes Ent. 1, p. 40.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1534 (see for synonymy, additional refer-
ences, and list of species).
Jedlicka 1963, Ent. Abhandlungen 28, pp. 489,
498.
Ophinnca Eschscholtz 1829, Zool. Atlas 2, p. 5
(not Ophionea Klug 1821).
Liebke 1938, Festschrift Embrik Strand 4, p. 79.
Diagnosis. See preceding Key to Genera.
Description. None required here.
Type species. Of both Casnoidea and
Ophionea Eschscholtz, Attelahus indica
Thunberg.
Generic distribution. SE. Asia including
Ceylon and Japan to Australia; a species
recorded also from the Seychelles Is.
Notes. This genus of slender, usually
strikingly bicolored (red and black) carabids
includes several species widely distributed
in SE. Asia and the Malay Archipelago.
They are usually found in grass, reeds, and
other vegetation growing in water.
Key to Species of Casnoidea of New Guinea
1. Color dark with basal Mi or -/-, of elytra red
(p. 207) ^estroi
- Color red with head, post-median elytral
fascia, and sometimes bases of elytra dark
(the post-median fascia with a pale spot on
each elytron) 2
2. Pronotum conspicuously punctate (p. 207)
ptiHcticollis
- Pronotum not conspicuously punctate 3
3. Elytra not dark at l)ase (p. 208) -.
( nigrofasciaia )
- Elytra dark at base (p. 208) _.... (imJira)
Casnoidea gesfroi (Maindron)
Maindron 1910, Bull. Soc. Ent. France for 1910,
p. 34 ( Oi)J}ionc(i) .
Dupuis 1913, Ann. Soc. Ent. Belgium 57, p. 270.
Liebke 1938, Festschrift Embrik Strand 4, p. 79,
fig. 60 {Ophionea).
gestiunis Seidlitz 1912, Archiv fiir Naturgeschichte
77, Part 3, p. 155 (error for gestroi).
Description. None required here; see
preceding Key to Species; length c. 7.0-
7.5 mm.
Type. From Dilo, Papua, July 1890 (D.
Loria); presumably in Paris Mas. (not
seen ) .
Occurrence in New Guinea. Papua: 7,
Dobodura, Mar.-July 1944 (Darlington); 2,
Kiunga, Fly R., July 15-21, Aug. 1-3, 1957
(W. W. Brandt, Bishop Mus.). West N. G.:
1, Waris, S. of Hollandia, 450-500 m, Aug.
8-15, 1959 (T. C. Maa, Bishop Mus.); 1,
Wasian, Vogelkop, Sept. 1939 (Wind,
M.C.Z.).
Notes. Tliis distinct species is evidently
widely distributed in New Guinea and is
represented also on New Britain (an un-
described subspecies from Cape Glouces-
ter) but is unknown elsewhere.
Casnoidea puncficollis (Sloane)
Sloane 1923, Proc. Linnean Soc. New South \\'ales
48, p. 31 (Ophionea).
Liebke 1938, Festschrift Embrik Strand 4, p. 80
(Ophionea).
Description ( for recognition only ) . Form
as in Figure 127; red, head black, elytra
with broad transverse fascia ( bluish ) black,
the fascia with an elongate pale fleck on
each 5th interval, legs bicolored; shining.
20S Bulletin Miiscmu of Comparative Zoology, Vol. 137, No. 1
without reticulate microsculpture. Head
short, rounded posteriorly; front wrinkled
anteriorly, impunctate. Prothorax long-
oval; side margins irregularly indicated
anteriorly; disc conspicuously punctate.
Eh/fra punctate-striate. Measurements (New
Guinean specimen): length c. 7.5; width
2.1 mm.
Type. From Burdekin R., Queensland,
Australia; in Sloane Coll., C.S.I.R.O., Can-
berra ( seen ) .
Occurrence in New Guinea. Papua: 1,
Kiunga, Fly R., Aug. 1-3, 1957 (W. W.
Brandt, Bishop Mus.).
Notes. I do not have puncticoUis from
Australia and have identified the New
Guinean individual from description. Note
that C. ^estroi (preceding species) as well
as puncticoUis occurred at Kiunga.
iCasnoidea nigrofasciafa (Schmidt-Goebel))
Schmidt-Goebel 1846, Faumila Coleop. Bir-
iiianiae, p. 21 (Opiiionea) .
Aiidrewes 1930, Treubia 7, Suppk-ineut, p. 3'34
( Ophionea ) .
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1535 ( see for synonymy and additional refer-
ences ) .
I.iebki' 1938, Festschrift Embrik Strand 4, p. 80,
fiji. 57 (Opiiiouca ).
Description. None required here; see
preceding Key to Sj)ecies.
Type. From Buruia; in Prague Mus.
(not seen).
Occurrence in New Guinea. Probably
does not occur.
Notes. C. ni<irofasciaia ranges from SE.
Asia to Java and Borneo. It is apparently
not recorded bom Celebes or the Moluccas.
New Guinea is included in the species'
range by Csiki, but I can find no authority
for this. 1 suspect that a too-hasty com-
piler, not noticing the negative, picked
"New Guinea" out of Andrewes' (1930)
statement that "I havi' seen no examples
cithci' hoin Jaj^an or New (iuinea."
(Casno/c/ea indica (Thunberg))
i'lmnbc'ru 1784, Novas Inseclorinn Species .3, p.
()8, flu. 81 (Altclalms).
Andrewes 1930, Cat. Indian Insects, Part 18,
Carabidae, p. 241 {Ophionea).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1534 ( see for synonymy and additional refer-
ences ) .
Licbke 1938, Festschrift Embrik Strand 4, p. 79,
ti^. 55 ( Ophionea).
Louwerens 1958, Treubia 24, p. 249 (Moluccas)
{Oi)]uonea).
Description. None required here; see
preceding Key to Species.
Type. From "India orientali"; presumed
lost (not seen).
Occurrence in New Guinea. Doubtful.
Notes. This common Oriental carabid
ranges from SE. Asia including Ceylon
and Japan to Celel>es and the Moluccas.
A specimen in the British Museum is labeled
"Dory, New Guinea" but may be from
Celebes or the Moluccas ( see Part I of m\
"Carabid Beetles of New Guinea," p. 331).
Other collectors have failed to find the
species in New Guinea. Andrewes' state-
ment that indica occurs south to New
Cruinea is probabh' based on the doubtful
"Dory specimen.
Genus BASISTICUS Sloane
Sloane 1917, Proc. Linnean Soc. New South Wales
42, p. 415.
1923, Proc. I.innean Soc. New South
Wales 48, p. 30.
Liebke 19.38, Festschrift Emlirik Strand 4, p. 81.
Diapiosis. See preceding Key to Genera.
Description. None required here.
Type species. Odacant}ia micans Macleay
( below ) .
Generic distribution. As of the single
known species.
Notes. This genus is close to CU)lliuris
(sensu Jafo ), from which it differs in luuing
the lateral margins of the prothora.x entire.
Basisficus micans (Macleay)
MacleaN 18()4, 'I'rans. Kwi. Soc. New South Wales
1, p. 107 (Oducanlha).
CsiVi 1932, Coleop. ('at., (Carabidae, Harpalinae 7,
p. 15.3.5 (see for additional references),
l.iebke 1938, Festschrift Fnibrik Strand 4, p. 81.
Description ( lor recognition ouK). Form
The Carabid Beetles of New Guinea
Darlinfitoii
209
of ColUuris; head and prothorax red, base of
elytra dark reddish, smooth part of elytra
piceous, antennae red, legs dark with pale
bases; elytra very coarsely punctate-striate
in anterior Vs, smooth with striae of minute
punctules in posterior %; length c. 6.5 mm.
Type. From Port Denison, northern Aus-
tralia ( presumably near Bowen, Queens-
land); probably in Macleay Mus., Sydney
(not seen).
Occurrence in New Guinea. Papua: 1,
Rouku, Morehead R., March 1962 (W. W.
Brandt, C.S.I.R.O.).
Notes. This is an Australian species, well
known in North Queensland. I have speci-
mens from the vicinity of Cairns, Mareeba,
and Townsville. The single individual from
New Guinea matches Australian ones well.
In Australia, this insect is found on the
ground in open woodland; the type was
"found under dried cow dung."
Genus CLARENCIA Sloane
Sloane 1917, Proc. Linnean Soc. New South Wales
42, p. 415.
1923, Proc. Linnean Soc. New South
Wales 48, p. 30.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1535.
Liebke 1938, Festschrift Embrik Strand 4, p. 81.
Diagnosis. See preceding Key to Genera;
note form of large CoUiuris; antennae with
very long 3rd segments (and see Notes,
below ) ; £ front tarsi with 3 segments with
numerous narrow squamae loosely ar-
ranged (not 2-seriate as in CoUiuris); last
ventral segment slightly emarginate at apex
in both sexes, with 1 seta each side in <^ , 2
in 9 .
Description. None required here.
Type .species. Casnonia aliena Pascoe, of
Australia.
Generic distribution. Eastern Australia,
New Guinea.
Notes. Although only 1 Clarencia is cur-
rently recognized (Csiki, 1932), 4 species
are represented in Australian material col-
lected by me in 1957-1958. One of these
species (described below as quadridens)
occurs also in New Guinea, and 1 additional
species of the genus is endemic in New
Guinea.
The antennae of some Australian Clar-
encia not only have very long 3rd seg-
ments but also have the 4th segments
uniquely modified: expanded and obliquely
truncate at apex so that the 5th segments
hinge forward, and with the pubescence
of the 4th segments restricted to the seg-
ments' anterior edges. This modification
of the 4th segments is only slightly indicated
in the New Guinean species, more clearly
in papua than in quadridens.
The species of Clarencia are usually
found in wet places, often by standing
water, either among wet leaves or in or
under low vegetation.
Key to Species of Clarencia of New Guinea
1. Elytra toothed at sutural and outer-apical
angles (p. 209) (iiiadridens
- Elytra not toothed (p. 210) \ni\>iia
Clarencia quadridens n. sp.
Description. With characters of genus;
form as in Figure 128, with elytra trans-
versely impressed near base; black, elytra ±
yellowish at apex but not spotted, epipleuri
pale, femora pale at base, dark at apex,
tibiae dark banded with pale, tarsi pale,
antennae brown darker basally; shining,
without distinct reticulate microsculpture.
Head 1.37 and 1.27 width prothorax; front
with conspicuous V-shaped impression an-
teriorly, impunctate. Prothorax long, with
sides swollen behind middle; width length
0.59 and 0.61; base apex 1.38 and 1.40;
disc very convex, with fine middle line,
punctate across base with a few punctures
along lateral margins and across apex but
otherwise impunctate. Elytra: width elytra/
prothorax 2.30 and 2.47; apices obliquely
sinuate-truncate with outer and sutural
angles acutely dentate; striae formed by
lines of punctures anteriorly, obsolete pos-
teriorly; 3rd intervals with c. 6 and 5th
intervals with c. 4 seta-bearing punc-
tures. Legs slender; tarsi above not pubes-
cent and not sulcate; 4th hind-tarsal seg-
210 Bulletin Museum of Comparative Zoology. Vol. 137, No. 1
ments shallovvly emarginate. Measurements:
length 10.0-11.5; width 2.8-3.1 mm.
Types. Holotype c^ (M.C.Z., Type No.
31,508) and 14 paratypes from Hollandia,
West N. G., Jiily-Sept. 1944 (Darhiigton);
and additional paratypes from West N. G.
as follows: 4, Hollandia, Apr., May 1945
(B. Malkin, U.S.N.M.); 2, "Neth. N. G."
without further locality (T. Aarons, Cal.
Acad.).
Additional material. West N. G.: 1
teneral S , Maffin Bay, Aug. 1944 ( Dar-
lington). Papua: 1 $, Normanby Is.,
Wakaiuna, Sewa Bay, Jan. 1-8, 1957 (W.
W. Brandt, Bishop Mus.).
Measured specimens. The S holotype and
1 9 para type from Hollandia.
Notes. The denticulate elytra distinguish
this species from all other known Clarencia
in Australia as well as New Guinea. This
species occurs in Australia: I have one
9 from near Cairns, N. Queensland (Dar-
lington). This Australian specimen and the
one from Normanby Is. have more yellow
at apex of elytra than Hollandia speci-
mens do.
Clarencia papua n. sp.
Description. With characters of genus;
form c. as in quadridens (above) except
elytra relatively narrower and not dentate;
])lack, elytra scarcely paler at apex, legs
bicolored as in quadridens, antennae brown;
shining, without distinct reticulate micro-
sculpture. Head 1.32 and 1.22 width pro-
thorax; front convex except impressed an-
teriorly as in quadridens, impunctate.
Frotliora.x formed as in (juadridens but
slightly shorter; width lengtli 0.62 and 0.68;
base/apex 1.37 and 1.41; disc very convex,
middle line fine, surface closely wrinkled-
punctate across base, variabK punctate
across apex, and more extensively punctate
at sides than in quadridens. Elytra: width
elytra prothorax 2.21 and 2.10; ajiices
oblicjuely sinuate-truncate with outer and
sutural angl(\s narrowK' rounded; striae
nearly entire (longer than in (piadridens),
punctate, the punctures becoming minute
posteriorly; 3rd and 5th intervals with a
few seta-bearing punctures. Legs c. as in
(juadridens. Measurements: length c. 9.0-
10.0; width 2.5-2.9 mm.
Types. Holotype $ (U.S.N.M.) and 7
paratypes (some in M.C.Z., Type No. 31,509)
from Hollandia, West N. G., Apr., May,
June (holotype, Apr.) 1945 (B. Malkin); 1
paratype, same locality, "11/5/44" (W. T.
Nailon, Fenton Coll.); 1 paratvpe, Yent-
chan. Main R., Sepik, N-E. N.' G., Feb.
1965 (R. Hornabrook).
Additional material. Papua: 2, Lake
Daviumbu, Fly R., Aug. 19-30, Sept. 11-20,
1936 (Archbold Exp., A.M.N.H.).
Measured specimens. The c5 holotype and
1 9 paratype.
Notes. See preceding Description for
details distinguishing papua from quad-
ridens. Actually, papua may be more closely
related to undescribed Australian species.
Genus DICRASPEDA Chaudoir
Chaudoir 1S62, Bull. Soc. Nat. Moscow 35, Part
2, p. 300.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1536 (see for partial s>n()n\in\ and addi-
tional references).
Liebke 1938, Festschrift Embrik Strand 4, pp. 43,
88.
Mdcroccntra Chaudoir 1869, Revue et Ma.uasin
Zool. (2) 21, p. 205 (new synonymy).
Liebke 1938, Festschrift Embrik Strand 4, pp. 39,
100.
Loxocdia Sloanc 1907, Dcutsihc Ent. Zeitscluitt
for 1907, pp. 179, 474.
Pliilcmonki Liebke 1938, Festschrift Enilirik
Strand 4, pp. 39, 83 (new s>n()n> ni\- ).
Diagnosis. Form as in Figiu'e 129 but
somewhat xariable. color black or metallic,
not maculate. Head: a line costa over each
eye; antennae with 3rd segments not or
not much longer than 4th segments. Pro-
tliora.x moderatelx long; pronotum chan-
neled at sides and with deep median
gr()()\('. Elytra: apici's \ariable ( sei' fol-
lowing Key to Species). Legs: tarsi not
jKibescent above (Liebke's statenunt that
tarsi ol Macroccntrd aic pubescent above
is erroneous); lib hiiid-tarsal segments
\ariab1y emarginate or lobed (see Notes,
The Carabid Beetles of New Guinea
Darlington
211
below). Secondary sexual characters: $
front tarsi narrow, narrowly 2-seriately
sqiiamulose; 6 with 1, 9 2 setae each side
last ventral segment.
Description. None required here.
Type species. Of Dicraspeda, D. brunnea
Chaudoir (see below). Of Macrocentra, M.
(juadrispinosa Chaudoir, of New Guinea.
Of Loxocara, L. quadrispinosa Sloane { =
M. quadrispinosa Chaudoir). Of PhiJemonia,
P. longiloba Liebke, of New Guinea.
Generic distribtition. Most diverse in
New Guinea; several New Guinean species
reach New Britain, etc.; species of Di-
craspeda sensu stricto (small forms with
unanned elytra) occur in Australia, and 1
(brunnea Chaudoir, below) extends to
Timor, Java, and the Philippines.
Notes. The 6 New Guinean species here
brought together in Dicraspeda are super-
ficially diverse, differing in form, presence
or absence of elytral spines, and form of 4th
hind-tarsal segments. But the differences
are all gradational (see following para-
graphs ) , different characters vary indepen-
dently, and the 6 species all share characters
given in the preceding Diagnosis. More-
over, they all inhabit understory foliage
of rain forest, and I think that they are all
probably derived from one ancestral stock
that has diversified in this habitat. Five
of the species are lowland forms and are
sympatric, occurring together at Dobodura.
The sixth species, D. ("Macrocentra")
violacea (Sloane), occurs at moderate alti-
tudes in the mountains.
The elytral apices are obliquely truncate
with sutural angles usually slightly blunted
and outer angles obtuse in D. (sensu
stricto) brunnea. In the 3 species of
"Philemonia" the sutural angles are either
slightly blunted (most individuals of longi-
Joba), variably denticulate (dubia and
some individuals of other species), or
spined (typical individuals of bispinosa);
the outer-apical angles are well formed in
these species and usually acute in longiloba,
but not spined. And in the 2 species of
"Macrocentra' (quadrispinosa and violacea),
outer-apical as well as sutural angles are
spined.
Variation of the 4th hind-tarsal segments
is noteworthy and is not correlated with
the insects' size or with form of elytral
apices. The 4th hind-tarsal segments are
shallowly emarginate (Fig. 163) in D.
(sensu stricto) brunnea; very deeply emar-
ginate with extremely long lobes (Fig. 165)
in D. ("PJiilemonia" ) longiloba, which re-
sembles brunnea in size and elytral apices;
and intermediate but variable in the other
species (other "Philemonia" and "Macro-
centra" ) .
Key to Species of Dicraspeda of New Guinea
1. Elytra without spines or with spines only at
sutural angles 2
- Elytra with spines at sutural and outer-apieal
angles 5
2. Fourth hind-tarsal segments emarginate for
c. Mi segments' length; elytra with sutural
angles slightly blunted, outer-apical angles
obtuse; length c. 5.5-6.0 mm (p. 211) - brunnea
- Fourth hind-tarsal segments more deeply
emarginate; elytra with sutural angles den-
ticulate or spined ( except in most longiloba ) ;
size larger 3
3. Fourth hind-tarsal segments very long-
lobed (Fig. 165); sutural angles usually
blunted; length c. 6.5-7.5 mm (p. 212) ..
longiloba
- Fourth hind-tarsal segments with shorter
lobes; sutural angles denticulate or spined;
size usually larger -— 4
4. Fourth hind-tarsal segments with lobes c.
^2 segments' length; sutural angles denticu-
late; length 6.5-8.0 mm (p. 212) ilulna
- Fourth hind-tarsal segments with longer
lobes; sutural angles spined or denticulate;
length 8.0-9.5 mm (p. 212) bi.s-pinosa
5. Color black; tarsi sulcate-carinate above;
length c. 11-13 mm (p. 213) quadrispinosa
- Color green-purple; tarsi not sulcate-carinate
above; length c. 11-12 mm (p. 213) -..
violacea
Dicraspeda brunnea Chaudoir
Chaudoir 1862, Bull. Soc. Nat. Moscow 35, Part 2,
p. 300.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1536 ( see for synonymy and additional refer-
ences ) .
Liebke 1938, Festschrift Embrik Strand 4, p. 89.
Description. None required here; note
212 BiiUctin Museum of Comparative Zoology. Vol. 137, No. 1
size small; pronotum punctate across base
and apex and in lateral and median grooves
but widely smooth at middle; elytral apices
unarmed; 4th hind-tarsal segments not
deeply emarginate (Fig. 163); length c.
5.5-6.0 mm.
Type. From Celebes; in Oberthiir Coll.,
Paris Mus. (not seen).
Occurrence in New Guinea. Papua: 9,
Dobodura, Mar.-July 1944 (Darlington);
1, Nonnanby Is., Wakaiuna, Sewa Bay, Jan.
1-8, 1957 (W. W. Brandt, Bishop Mus.).
N-E. N. G.: 1, Wareo, Finschhafen (Rev.
L. Wagner, S. Australian Mus.). West
N. G.: 3, Hollandia and vicinity (various
dates and collectors ) .
Notes. D. bninnea is recorded from
Australia (Queensland), Timor, Celebes,
Java, and Mindanao, and I have specimens
from Leyte and Luzon and New Britain
as well as from New Guinea.
Dicraspeda longiloba (Liebke)
Liehke 1938, Festschrift Einbrik Strand 4, p. 83
( F/i ilemonia ) .
Description. None required here; note
elytra with sutural angles blunt or at most
minutely denticulate; 4th hind-tarsal seg-
ments very long-lobed ( Fig. 165 ) ; length c.
6.5-7.5 mm.
Type. From N-E. N. G. ("Deutsch-Neu-
Guinea"); in Liebke Coll., present location
unknown (not seen).
Occurrence in New Guinea. N-E. N. G.:
the type. Papua: 5, Dobodura, Mar.-July
1944 (Darlington).
Notes. I have a specimen also from Cape
Gloucester, New Britain.
Dicraspeda dubia (Gestro)
(Jestro 1879, Ann. Mus. Civ. C.cnoa 14, p. 558
(Odacanlha) .
Liebke 1938, Festsebrift i;ii.l)rik Strand 4, p. 83,
fJLC. f:)7 {Fluh'Dionid) .
Descri])tion. None required here; note
elytra with sutural angles variably denticu-
late but not si)ined; 4th hiud-taisal seg-
ments rather short-lobcd but somewhat
varia])le; length c. 6.5-8.0 mm.
Type. From Fly R., presumably Papua;
in Genoa Mus. (not seen).
Occurrence in New Guinea. Papua: 3,
Dobodura, Mar.-Julv 1944 (Darlington);
5, Kokoda, 1200, 1300 ft.. May, Aug., Sept.,
Oct. 1933 (Cheesman). N-E. N. G.: 1,
Aitape, Aug. 1944 (Darlington); 1, Wau,
Morobe Dist., 1200 m, Aug. 18, 1961 ( Sed-
lacek), in light trap. West N. G.: 1, Waris,
S. of Hollandia, 450-500 m, Aug. 16-23,
1959 (T. C. Maa, Bishop Mus.); 1, Xabire,
S. Geelvink Bay, 0-30 m, July 2-9, 1962
(Gressitt); 1, Waigeu Is., Camp 1, Mt. Nok,
2500 ft. ( c. 760 m ), May 1938 ( Cheesman ) .
Notes. I found this or a closely related
species also at Bamaga, near the tip of
Cape York, Australia.
Dicraspeda bispinosa n. sp.
Description. With characters of genus;
form as in Figure 129; brownish black, legs
dark, antennae and mouthparts paler
brown; moderately shining, reticulate mi-
crosculpture indistinct on head and pro-
notum, isodiametric or slightK' transverse
on elytra. Head 1.17 and 1.17 width pro-
thorax; front irregular!)' impressed anteri-
orly, impunctate; mentum w ith long narrow
tooth; ligula 4-setose. Prothorax elongate-
subquadrate with sides swollen below mar-
gins; width length 0.99 and 0.98; base '
apex 1.19 and 1.12; pronotum strongly con-
vex, narrowly channeled each side near
margin and with well impressed middle
groove; surface punctate chiefly across
base, slightly at apex. Elytra: width elytra/
prothorax 2.08 and 2.05; apices obliquely
sinuate-truncate with sutural angles spiued
or denticulate, outer-apical angles obtuse,
and apical margin in part minuteh' d(>nticu-
late; striae entire, punctate; 3rd inter\als
3-punctate, the posterior puncture near
apex. Inner n'i)iij.s full. /.ri,'.v normal; tarsi
not suleate and not pubescent aboM'; 4th
hind-taisal segments long-lobed (Fig. 164).
Secondary sexual cJiaracters as of genus;
last Ncntral segment with small notch at
apex in both sexes. Mcdsiircinciils: length
The Carabid Beetles of New Guinea • Darlington 213
(including spines) c. 8.0-9.5; width 2.8-
3.3 mm.
Types. Holotype $ (M.C.Z., Type No.
31,510) and 15 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. Papua: 2,
Kokoda (Cheesman); 2, Mt. Lamington,
1300-1500 ft. (c. 400-460 m) (C. T.
McNamara, S. Australian Mus.). N-E.
N. G.: 4, Wau, Morobe Dist., 1150, 1200,
1300 m, dates in Jan., Feb., Oct., 1961, 1963
(Sedlacek); 1, Finschhafen, Huon Pen., 80-
200 m, Apr. 13, 1963 (Sedlacek).
Additional material. N-E. N. G.: 4,
Finschhafen, 80 m, Apr. 16, 1963 (Sedlacek);
1, same locality, 80-200 m, Apr. 13, 1963
(Sedlacek); 3, Pindiu, Huon Pen., dates
in Apr. 1963 (Sedlacek).
Mea.sured specimens. The 6 paratype and
9 holotype from Dobodura.
Notes. This new species would go in
Philemonia in Liebke's classification. The
specimens listed under Additional material
have the sutural angles of the elytra den-
ticulate rather than spined, but I think they
are referable to bispinosa. Note that both
spined and denticulate forms have been
found at Finschhafen.
Dicraspeda quadrispinosa (Chaudoir)
Chaudoir 1869, Revue et Magasin Zool. (2) 21, p.
206 ( Macrocentra ) .
Sloane 1907, Deutsche Ent. Zeitschrift lor 1907, p.
474 (Macrocentra).
Liebke 1938, Festschrift Embrik Strand 4, p. 100
( Macrocentra ) .
Louwerens 1956, Trculiia 23, p. 223 (Moluccas)
(Macrocentra).
Loxflcara quadrispinosa Sloane 1907, Deutsche Ent.
Zeitschrift for 1907, p. 180.
Description. None required here; known
among New Guinean Colliurini by size
large; color plain black; elytra spined at
sutural and outer-apical angles; tarsi sul-
cate-carinate above; length (including
spines) c. 11-13 mm (rarely shghtly smaller
or larger).
Types. Of quadrispinosa Chaudoir, from
Dorey, West N. G. ( \\'allace ) ; in Oberthiir
Coll., Paris Mus. Of quadri.spinosa Sloane,
from Simbang, N-E. N. G.; "returned to
Dr. Horn (for Bennigsen's collection)"
(not seen).
Occurrence in New Guinea. Common:
227 specimens from localities on New
Guinea and Normanby, Ferguson, Wood-
lark, Rossel, Sudest, and Waigeu Is.; ap-
parently confined to low altitudes, up to
700, 750, 800, and 975 m at different
localities, but none found higher; cnmmnn
at Dobodura.
Notes. This characteristic New Guinean
carabid has been foimd also in the Moluc-
cas, New Britain, and the Solomons, but
not Australia.
Dicraspeda violacea (Sloane)
Sloane 1907, Deutsche Ent. Zeitschrift for 1907,
pp. 181, 474 (Macrocentra).
Liebke 1938, Festschrift Embrik Strand 4, p. 100
(Macrocentra).
hahiJis Sloane 1907, Deutsche Ent. Zeitschrift for
1907, p. 181 (name used in error for violacea).
Description. None required here; similar
to preceding (iptadrispinosa) but head and
pronotum greenish or bluish, elytra purple;
tarsi not sulcate-carinate above; length c.
11-12 mm.
Type. From Sattelberg, N-E. N. G.; "re-
turned to Dr. Horn (for Bennigsen's col-
lection)" (not seen).
Occurrence in New Guinea. N-E. N. G.:
8, Wau, Morobe Dist., 1200, 1300 m, Jan.,
Mar., Apr., June, Sept., Nov., 1961-1963
(Sedlaceks); 1, Eliptamin Vy., 1665-2530
m, June 23-30, 1959 (W. W. Brandt, Bishop
Mus.); 1, Finisterre Rge., Saidor, Kiambavi
Village, July 22-29, 1958 (\^^ W. Brandt,
Bishop Mus.); 1, W^areo, Finschhafen (Rev.
L. Wagner, S. Australian Mus.); 1, Goroka,
E. Highlands, 5200 ft. (c. 1600 m), J. H.
Barrett, Dept. Agr. Port Moresby). West
N. G.: 3, Rattan Camp, Snow Mts., 1150,
1200 m, Feb.-Mar. 19.39 (Toxopeus); 5,
Fac Fac, Vogelkop, 100-700 m, June 9,
1959 (Gressitt and T. C. Maa, Bishop
Mus.), in light trap; 1, Mt. Baduri, Japen
Is., 1000 ft., Aug. 1938 (Cheesman).
Notes. D. violacea apparently replaces
quadrispinosa above 1000 m altitude in
214
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
New Guinea, Init the two species overlap
below 1000 m. D. violacea occurs also in
New Britain (Gaulim, Gazelle Pen., 130 m,
Nov. 28, 1962, Sedlacek).
Genus LACHNOTHORAX Motschulsky
Motsthulsky 1862, fitude Ent. 11, p. 48.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1542 (see for additional references).
Liebke 1938, Festschrift Embrik Strand 4, p. 103.
Jeannel 1948, Coleop. Carabiques de la Region
Malgache, Part 2, p. 7.56.
Diaii,nos-is. See Key to Genera of Col-
liurini of New Guinea; this is the only con-
spicuously pubescent colliurine in New
Guinea.
Description. None required here.
Type species. L. higuttatus Motschulsky,
of India.
Generic disfrihtition. Africa and Mada-
gascar; SE. Asia to New Guinea.
Notes. The few species of Lachnothorax
are all very much alike. They are probably
ground-living, and I suspect that they occur
by running water.
Lachnothorax. tokkia Gestro
Gestro 1875, Ann. Mns. Civ. Cenoa 7, p. 856.
Csiki 1932, Coleop. ('at., Carabidae, Harpalinae 7,
p. 1542 (see for synonymy and additional refer-
ences ) .
Liebke 1938, Festschrift Embrik Strand 4, p. 101.
Jedlicka 1963, Ent. Abliandlnngen 28, p. 504.
Description. None required here; note
form, color black with pale spot before
apex each elytron, and consjiicuous pubes-
cence; length c. 5.0-5.5 mm.
Type. From Kandari, SE. Celebes; in
Genoa Mus. (not seen).
Occurrence in New Guinea. N-E. N. (i.:
7, Stephansort, Astrolabe Bay, 1900 (Biro).
Notes. Lachnothorax tokkia has been r(^-
corded previously from the Malay Pen..
Sumatra, Java, and Celebes, and a ver\
closely relat(>d species {hi<iuttata Motschul-
sky) occurs in India and Geylon.
Genus EUDALIA Castelnau
Caslelnan 1867, Notes on Australian Coleop., p. 16.
Sloane 1917, Proc. Linnean Soc. New South Wales
42, pp. 415, 417-422 (with key to Australian
species ) .
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1542 (see for synonymy and additional refer-
ences ) .
Liebke 1938, Festschrift Embrik Strand 4, pp. 44,
105.
Diapiosis. See Key to Genera.
Description. None required here.
Type species. Odacantha latipennis Mac-
leay, of Australia.
Generic distribution. Australia (c. 6
species ) and New Guinea ( 1 species,
doubtfully assigned to this genus).
Notes. The species described below as
Eudalia anomala has entire lateral pro-
thoracic margins and therefore runs to
Eudalia in Liebke's key, but if the margins
were obsolete, it would run to Andrewesia,
to which it may be more closely related.
{Andrewesia ohcsa (Andrewes) ranges
from the Malay Pen. to the Moluccas.)
Generic characters and limits in this group
(as in so many others!) need revision.
Eudalia anomala n. sp.
Description. Form as in Figure 130;
black, elytra faintly aeneous and tipped
with yellow, legs testaceous, antennae
brown paler at base; h(>ad and pronotum
shining and without reticulate microsculp-
ture, elytra duller with isodiametric meshes.
Head 1.18 and 1.17 width prothorax;
strongly constricted at neck; antennae with
segments 3 and 4 sube(jual, pubescent from
4th segmcMits; mandibles moderate^ in knigth
and curvature; front eon\ex, irregularly
impressed anteriorK', impunetate; mentuin
with moderate tooth; liguJa broad, with 2
long and 2 shorter setae; palpi slender, not
pubescent. Protliorax suborbicular except
parallel at base; width length 0.89 and
0.90; base apex 1.25 and 1.26 (sides of pro-
thorax curve into condyle of neck at apex);
lateral margins narrow but (Mitire, with a
seta-bearing pimcture inside margin (on
disc) before middle; disc \('ry convex,
strongly transverseh- impressed at base;
middle line slightly impressed; surface
The Carabid Beetles of New Guinea • Daiiinaiott
215
punctate across base, impunctate or nearly
so elsewhere. Elytra ample; width elytra,
prothorax 2.15 and 2.16; apices obliquely
truncate with outer angles obtuse and
inner angles acute-blunted; striae entire,
punctate; 3rd intervals with 4 or 5 seta-
bearing punctures including 1 near base.
Inner winp,s full. Legs moderate; tarsi not
pubescent above and not sulcate; 4th hind-
tarsal segments emarginate but not lobed.
Secondary sexual characters: i front tarsi
narrow, 2-seriately squamulose; last ventral
slightly emarginate at apex in 6 , not in $ ,
with 1 seta each side in i , 2 in 9 . Mea-
surements: length c. 7.0; width 2.5 mm.
Types. Holotype $ (A.M.N.H.) and 1
i paratype (M.C.Z., Type No. 31,511)
from Menapi, Cape Vogel Pen., Papua,
0-30 m, "No. 1," Aug. 8-11, 1953 (Geoffrey
M. Tate); and 1 9 i^aratype from Wasian,
Vogelkop, West N. G., Sept. 1939 (Wind,
M.C.Z.).
Measured specimens. The i holotype and
9 paratype.
Notes. For possible relationships of this
species, see under genus. This species is
smaller and much less punctate and less
roughened above than any typical (Aus-
tralian) Eudalia known to me, and the
New Guinean species has the outer-apical
elytral angles more angulate.
DOBODURA n. gen.
Diagnosis. See Key to Genera of Col-
liurini of Netv Guinea.
Description. Form ( Fig. 131 ) c. as in
some CoUiuris. Head without supraocular
costae; mandibles long, slender, weakly
arcuate; antennae very long, 3rd segments
c. V4 longer than 4th segments, 1st segments
with 1 long seta near apex; mentum with
triangular tooth; ligula rounded, with 2 long
setae at apex and 2 shorter setae laterally;
paraglossae small, membranous; palpi slen-
der, not pubescent. Prothorax: lateral mar-
gins entire; median impressed line fine;
base deeply transversely channeled with
transverse ridge behind channel. Elytra
spined. Inner icings full. Legs slender;
tarsi not pubescent above, not sulcate
above; 4th hind-tarsal segments moderately
emarginate, emargination c. '/•; length of
segment; claws not toothed. Secondary
sexual characters: 6 front tarsi scarcely
dilated, 3 segments narrowly 2-seriately
squamulose; last ventral segment of $
weakly, of 9 subcircularly emarginate,
with 1 seta each side in i , 2 in 9 .
Type species. D. armata (below).
Generic distribution. The single species
is known only from New Guinea.
Notes. This striking new genus may be
related to Eudalia but differs in form,
longer and less arcuate mandibles, much
longer antennae with relatively longer 3rd
segments, and presence of elytral spines.
The position of the principal spines, c.
opposite the ends of the 4th intervals
rather than at the sutural or outer-apical
angles, is unusual in this tribe.
Dobodura armata n. sp.
Description. With characters of genus;
form as in Figure 131; black above and
below, appendages testaceous except femora
dark on inner sides; shining, reticulate
microsculpture absent or indistinct on most
of upper surface, present and c. isodiametric
on elytra posteriorly. Head 1.07 and 1.06
width prothorax; front evenly convex except
slightly impressed anteriorly, impunctate;
neck slightly constricted. Prothorax suboval,
swollen at sides below margins; width
length 0.92 and 0.91; base/apex 1.14 and
1.15; margins each with seta-bearing punc-
ture c. -o from apex; disc c. evenly convex,
impunctate. Elytra: width elytra prothorax
2.00 and 2.09; apices with sutural and outer
angles denticulate or short-spined and with
long spines c. opposite ends 4th intervals;
striae entire, formed by lines of fine punc-
tures; 3rd intervals with 3 well spaced
seta-bearing punctures. Secondary sexual
characters as of genus; S copulatory organs
as in Figure 180. Measurements (types);
length (including spines) c. 10.5-11.5;
width 3.3-3.5 mm.
216 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Types. Holotype S (M.C.Z., Type No.
31,512) and 5 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
2 paratvpes, Mt. Hansemann, Astrolal^e
Bay, N-E. N. G. (Biro).
Additional material. West N. G.: 1 $ ,
mountain slope above Bernhard Camp, 100
m, Apr. 8, 19.39 (Toxopeus).
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. My specimens were taken among
spray-drenched stones beside small torrents
in rain forest.
The specimen from Bernhard Camp has
the strial punctures of the elytra coarser
than in the types and the tip of the aedeagus
slightly different. Additional material may
sho\\' it to represent a distinguishable sub-
species.
Tribe DRYPTiNI
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1548 (see for synonymy and additional refer-
ences ) .
Jeannel 1949, Coleop. Carabicjues de le Region
Malgache, Part 3, p. 1063.
Jedlicka 1963, Ent. Abhandlunyen 2S, p. 4S1.
Halm 1967, Fanna Jap<"iica, Caral)idae, Trun-
eatipennes Group, p. 266.
Drypfiddc Jeannel 1942, Faune de France, Coleop.
Carabicjues, Part 2, p. 1098.
Dryptimie Basilewsky 19.53, Exploration Pare Na-
tional rUpemba, Ease. 10, Carabidae, p. 228.
Members of this small but widely distrib-
uted tribe (represented in New Cuinea
by only 2 genera) are easily recognized by
characteristic form (P'ig. 132); pubescent
surface; antennae with very long 1st and
very short 2nd segments; and elytra with-
out raised outer margins. The New Guinean
species live chiefly in grass, 1 think. They
are winged and probably diurnal.
Key to (;kni:ha of Dkyi'tim ok New Guinea
I. CHaws simple (p. 216) _ /);■(//)/«
- Claws pectinate (p. 218) Dcscra
Genus DRYPTA Latreille
Latreille 1796, Precis Caracteres Generiqnes In-
sectcs, p. 75.
Csiki 1932, Coleop. (]at., Carabidae, Harpalinae 7,
p. 1.548 (see lor additional reterences).
Andrewes 1936, Proc. R. Ent. Soc. London (B)
5, p. 134 (key to "Indian" species).
See also references under tribe.
Diaiinosis. See characters given for tribe
and in preceding Key to Genera.
Description. None required here.
Type species. Carahus dentatus Rossi, of
Europe, etc.
Generic distribution. Tropical and warm
temperate regions of the Old World; 1
species listed from Brazil.
Notes. The Oriental-Australian species of
Drypta are much alike, differing chiefly in
proportions and color.
Key to Species of Drypta of New Guinea
1. Head, prothorax, and elytra blue-green (p.
216 ) fkipua
- Head and prothorax red; elytra brown,
black, blue-black, or striped 2
2. Elytra broadly longitudinally striped with
red (p. 217) _ inaatcrsi
- Elytra not striped 3
3. Less slender ( prothoraeic width dength r.
0.78 or more); femora dark (p. 217) fumi^ata
- More slender ( prothoraeic width length c.
0.7.5 or less); femora pale (p. 217) siilcicollis
Drypta popua n. sp.
Description. With characters of genus;
form as in Figure 132, c. average in genus;
greenish blue, appendages yellow with
apices of femora and of 1st antennal seg-
ments narrowly darker; entire upper sur-
face closely punctate. Head 1.09 width
prothorax; eyes moderate, genae convex.
Protliora.x subcylindric; width length 0.76;
base apex 1.08; lateral margins indistinct;
middle line poorly defined. Elytra: width
elytra prothorax 2.09; apices obliciueK sub-
truncate with outer angles obtuse-blunted
and sutmal angles c. right; striae impressed,
coarsely ver\ closely punctate; inten^als
more finely, less closely punctate. Le^s:
tarsi not sulcate abo\(>; 4th hiud-tarsal seg-
ments long-lobed; claws cm\(>d, not pecti-
nate, each w ith obtuse angulation of innc^r
edge near base but with no trace of teeth.
Secondary sexual characters not determined
{a unknown). Measurements: length 8.5;
w idth 2.7 mm.
Type. Holotype 9 (M.C.Z., Type No.
The Carabid Beetles of New Guinea
Darlington
217
31,513) from Lae, N-E. N. G., Oct. 1944
(Darlington); the type is unique.
Notes. I am not sure of the relationships
of this unexpected species. In Andrewes'
( 1936 ) key to "Indian" species of Drijpta
(see reference under genus), papua runs
to couplet LS ( 19 ) but fits neither species
there named, having a relatively narrower
head and broader prothorax than aeiheria
Andrewes (of Assam) and more closely
punctate elytral intervals than cijanopa An-
drewes (of Bengal). Drypta papua does
not resemble any Australian species of the
genus. It does superficially resemble
Desera cleg,ans Sloane (below) but is
smaller, with relatively broader prothorax
and obtuse rather than acute outer-apical
elytral angles, and of course with simple
rather than pectinate tarsal claws.
Drypta mastersi Macleay
Macleay 1871, Trans. Ent. Soc. New South Wales
2, p. 82.
Chaudoir 1877, Bull. Soc. Nat. Moscow 52, Part 1,
p. 257.
Description. None rec^uired here; note
elytra striped with red; length (of New
Guinean specimen) c. S.5 mm.
Type. From Gayndah, South Queensland,
Australia; presumably in Macleay Mus.,
Sydney (not seen).
Occurrence in New Guinea. Papua: 1,
Rouku, Morehead R., W. Papua (opposite
the tip of Cape York), Apr. 1962 (W. W.
Brandt, C.S.I.R.O.).
Notes. In Australia, mastersi ranges north
at least to mid-peninsular Cape York. I
do not know whether it is really distinct
from Drypta australis Dejean of more-south-
ern Australia.
Drypta fumigata Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 720.
Chaudoir 1877, Bull. Soc. Nat. Moscow 52, Part
1, p. 258.
Description. None required here; length
13.5-15.0 mm.
Type. From Andai, Papua, Aug. 1872
(Beccari and D'Albertis); in Genoa Mus.
(not seen).
Occurrence in New Guinea. Papua: 3,
Dobodura, Mar.-July 1944 (Darlington); 5,
Kiunga, Fly R., dates from Julv 23 to
Sept. 25, 1957 (W. W. Brandt,' Bishop
Mus.); 1, Owen Stanley Rge., Goilala,
Rome, 1950 m, Apr. 1-15, 1958 (W. W.
Brandt, Bishop Mus.); 1, Popondetta, N.
Dist., Jan. 29, 1965 (R. Hornabrook). N-E.
N. G.: 1, Erima, Astrolabe Bay, 1896
(Biro); 1, Chimbu Vv., Bismarck Rge.,
5000-7500 ft. (c. 1500-2300 m), Oct. 1944
(Darlington); 1, Aiyura, E. Highlands,
5600 ft. (c. 1700 m), "9.10.1960" (J. H.
Barrett, Dept. Agr. Port Moresby), at light;
2, Okapa, June 23, 1965 (R. Hornabrook).
West N. G.: 2, Hollandia, Apr., May 1945
(R. Malkin, U.S.N.M.); 32, Sansapor, Aug.
1944 (Darlington).
Notes. This species is presumably of
Oriental origin, but I cannot determine to
which Oriental species it is most closely
related.
Drypta suicicollis Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 721.
Chaudoir 1877, Bull. Soc. Nat. Moscow 52, Part 1,
p. 258.
Description. None required here; length
c. 10.5-11.5 mm.
Type. From Andai, Papua, Aug. 1872
( Reccari and D'Albertis ) ( note locality
same as for type of fumigata); in Genoa
Mus. (not seen).
Occurrence in New Guinea. Papua: 1,
Dobodura, Mar.-Julv 1944 (Darlington);
3, Kiunga, Fly R., Aug. 1-3, 14-17, 1957
(W. W. Rrandt, Bishop Mus.). N-E. N. G.:
1, Erima, Astrolabe Bay, 1897 (Biro); 1,
Stephansort, Astrolabe Bay, 1897 (Biro).
West N. G.: 1, Hollandia-Binnen, 25 m,
Oct. 16, 1957 (Gressitt); 1, Humboldt Bay
Dist., 1934 (British Mus.); 3, Tor R.
(mouth), 4 km E. of Hoi Maffen, July 2,
1959 (T. C. Maa, Rishop Mus.). at light;
2, Wasian, Vogelkop, Sept. 1939 (Wind,
M.C.Z.).
Notes. This species too is probably de-
rived from an Oriental (not Australian)
stock, but I do not know its exact relation-
ships.
218 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Genus DESERA Hope
Hope 1831, Zoological Mistellany, p. 21.
Csiki 1932, Coleop. Cat., Carabidae, Haipalinae 7,
p. 1553 (see for additional references).
Andrewes 1936, Proc. R. Ent. Soc. London ( B )
5, p. 136 ( kev to "Indian" species).
1939, Ann. Mag. Nat. Hist. (11) 3, p. 133.
Dendrocelhis Schmidt-Goebel 1846, Faunula
Coleop. Birmaniae, p. 24.
Diafinosis. Characters as of Drijpta, ex-
cept cla\\'s pectinate.
Description. None required here.
Ty))e .species. Dcsera nepalen.si.s Hope,
of SE. Asia (see following Note.s).
Generic distribution. SE. Asia to Aus-
tralia; Africa.
Note.s. Dcsera differs from Drypto ap-
parently only in having pectinate tarsal
claws. A modern revision of the species is
needed to show whether both genera are
really nionophyletic and distinct.
Andrewes ( 1939 ) outlines the history of
the name Dcsera. It was used by Hope
( 1S31 ) in combination with the valid de-
scription of a new species ( )icpalcnsis,
which is therefore the type species), and
the combined description iricludes refer-
ence to the pectinate tarsal claws. This
use validates Dcsera Hope 1831 under
Article 16(a) (VI) of the 1964 edition of
the International Code of Zoological No-
menclature.
A single, common species ol this genus
occurs in New Cuinea.
Desera elegans (Sloane)
Sloanc 1907, Dcutsilic Knt. Zcitsclirilt lor 1907,
p. 473 ( Dcii(lrocclliis).
Andrewes 1927, Ann. Mag. Nat. Hist. (9) 19, p.
110.
Description ( for recognition only). With
characters of tribe and genus; form slender;
green (sometimes l^lue-green or l)roiv/e-
green), antennae red with 1st segment dark
at apex, legs red with knees usualK' darker
(legs and antennae sometimes more ex-
tensively dark); knigth c. 9.5-10.5 mm.
Ty])C. Froiu Haining Berge, Cazelle Pen.,
New Itritaiii; in Deutsche Ent. Instituti',
Berhn-Dahlem (Andrewes 1927) (not seen).
Occurrence in New Guinea. Common:
197 specimens from many localities widely
scattered over New Guinea, from sea level
to c. 1700 m altitude; occurs at Dobodura
and \\'au.
Notes. Desera clciians of New Guinea,
New Britain, and New Ireland is similar
to genicidata King (SE. Asia to the Moluc-
cas) on one side and to smaraiidula
Chaudoir (Australia) on the other. In fact
a single individual from Rouku, Morehead
R., West Papua, Apr. 1962 ( W. W. Brandt,
C.S.I.R.O. ) looks more like the Australian
smarag,dula than like the New Guinean
elegans. Relationships (or identities?) of
these and other similar species in the
whole Asiatic-Australian area need further
study.
Tribe ZUPHIINI
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1562 (see for synonymy and additional refer-
ences ) .
Habn 1967, Fanna Japonica, Carabidae, Tnm-
catipennes Gronp, p. 253.
Ztiphiidae Jeannel 1942, Fanne de France, Coleop.
Carabiqnes Part 2, p. 1091.
ZitpJiiitac Jeannel 1949, Coleop. Carabiciues de la
Region Malgaelie, Part 3, p. 1047.
Ziiphiiuae Basilewsky 1953, Fxploration Pare Na-
tional I'l'pcniba I'^ase. 10, p. 224.
This is another small but wideK distrib-
uted tribe. Its characters and taxonomic
liiuits need not be discussed here. It is
represented in New Cuinea by 2 easily
recognized genera and a total ol 6 known
species.
The members of the tribe \\\r in wt't
places, often among dead lea\cs on tlu'
ground {Yjipliiuni) or in gniss and vegeta-
tion growing in water (Planctes). Most
sjieeies, including all those in New Cluinea,
are winged.
K\:\ ro Ckmvuv oi /.ii'iium oi Niw Guinea
1. Head Mil)triangiilar, \cr\ wide at base: eKtra
not eostate (p. 219) Zujiliitnii
- Head normal; elytra wilii Mian> line (.ostac
(p. 220) '- - I'laiictcs
The Carabid Beetles of New Guinea • Darlington 219
Genus ZUPHIUM Latreille
Latreille 1806, Genera Crustaceorum et Iiisectonim
1, p. 198.
Csiki 1932, Coleop. Cat., Caiabidae, Harpalinae 7,
p. 1562 (see for synonymy and additional refer-
ences ) .
Jedlicka 1963, Ent. Abhandlungen 28, p. 477.
See also references under tribe.
Diagnosis. Form (Fig. 133) diagnostic;
and see preceding Key to Genera.
Description. None required here.
Type species. Carahus oJens Rossi, of
Europe, etc.
Generic distribution. All tropical and
some temperate regions of the world.
Notes. The various Oriental and Austra-
lian species of Zuphium are not well under-
stood. For example, the 7 listed Australian
species were all described between 1867
and ISScS, most of them from single speci-
mens or single localities, and they have
never been revised. The real relationships
of the 2 New Guinean species are therefore
doubtful, although I have made some com-
parisons.
Besides the 2 species recorded from New
Guinea below, I have seen (British Mus.)
1 specimen of Zuphium celehense Chaudoir
labeled as from Dory, presumably collected
by Wallace. I think this specimen is prob-
ably really from Celebes ( see Part I of my
"Garabid Beetles of New Guinea," pp. 330-
331 ) , and I see no reason to list the species
from New Guinea even tentatively.
Key to Species of Zuphium of Ne\v Guinea
1. Large (c. 8.3 mm); color piceous (p.
219 ) thouzeti
- Small (c. 3.5 mm); color brown (p. 219)
simiinu
Zuphium thouzeti Castelnau
Castelnau 1867, Notes on Australian Coleop., p. 17.
1868, Trans. R. Soc. Victoria 8, p. 103.
Description. None required here. Note
size large; color dark, not spotted; length
(of New Guinean specimen) c. 8.3 mm.
Types. From Rockhampton, Queensland,
Australia; present location of type un-
known (not seen).
Occurrence in New Guinea. Papua: 1
9, Port Moresby, Feb.-May 1943 {W. B.
Jones, U.S.N.M.).
Notes. Besides the types from Rock-
hampton, Gastelnau had a specimen from
Port Denison (near Bowen) farther north,
and I have specimens (identified from de-
scription) from W. of Ravenshoe and N. of
Mareeba still farther north in Queensland.
The Port Moresby specimen agrees rea-
sonably well with my Australian ones ex-
cept that the color of the legs varies.
Zuphium sinuum n. sp.
Description. Form as in Figure 133, very
small; brown, head slightly darker, append-
ages and lower surface paler; dull, entire
upper surface densely microreticulate or
roughened. Head 0.92 and 0.94 width pro-
thorax; antennae short, middle segments c.
2x long as wide; surface densely micro-
reticulate, moderately punctulate. Pro-
thorax: width/length 1.13 and 1.10; base/
apex 0.88 and 0.81; posterior angles right-
acute and not quite basal ( base very briefly
subpedunculate); surface closely roughened-
punctate. Elytra: width elytra prothorax
1.65 and 1.70; apices sinuate at middle of
width, lobed between sinuations and suture;
surface roughened, striae indicated but not
well defined. Secondary sexual characters:
i front tarsi slightly dilated, 3 segments
with soles of dense short squamae; c^ with
1, 9 2 setae each side last ventral segment.
Measurements: length c. 3.5; width 1.3-
1.4 mm.
Types. Holotype 9 (M.G.Z., Type No.
31,514) from Aitape, N-E. N. G., Aug. 1944
(Darlington); and 1 6 paratype, Kota
Nika, Res. Hollandia, West N. G., Jan. 31,
1956 (R. T. Simon Thomas, Louwerens
GolL).
Measured specimens. The S paratype and
9 holotype, in this order.
Notes. This species or a close relative
occurs also at Cape Gloucester, New Brit-
ain. Small size, color, dull surface, and
sinuate elytral apices distinguish sinuum
from other comparable species including
220 BuUc'tin Mu.scin)i of Comparative Zoolog,!), Vol. 137, No. 1
cclc'])cnsc Chaudoir (see under genus), in
whieh the elytral apices are scarcely sinuate.
Z. inconspicuum Schmidt-Goebel of Burma,
etc., has strongly sinuate elytral apices but
is much more shining than .sintiinn.
Genus PLANETES Macleay
Macleay 1825, Aniuilosa Javanica, p. 28.
Csiki 1932, Coleop. Cat., Caiahidae, Harpalinat- 7,
p. 1567 (see for syiuiiiyiiiy and aclclitional refer-
ences).
Jcdiieka 1963, iMit. Ahhandlnnuen 28, p. 464.
Di(ii!,n()si.s. Form c. as in Figure 134;
elytra each with more than 20 line longitu-
dinal costae.
Description. None required here.
Type species. P. himacuJatus Macleay,
of Java, etc.
Generic distrihution. SE. Asia including
Oylon and Japan to northern Australia;
Africa.
Notes. Most and most diverse species of
Planetes occin- in the Oriental Region. Four
species, all rather small and imspotted, are
known in New Guinea. And only I or 2
species, the same as or close to New
Guinean lorms, ha\t' l)een found in Aus-
tralia. This geographic pattern suggests
dispersal from Asia to Australia.
The species of this genus that I have
collected live in swamps and beside stand-
ing water. The\' are \\inged and often fly to
light.
In the present work I have not distin-
guished Uetero^Jossa Nietner from Planetes,
although the two probably are distinct (see
Ihibu, 1967, reference cited under tribe).
Ki;v K) SiM'CiKs OF Planetes ok Nkw CIuinka
1. Smaller, lenijtli 6.0—7.5 mm; see also l^c-
■srrii)li()u (p. 220) scccnuiutus
— Usually larger; // leniilli under 8 mm, 19th
( posthiimeral ) elytral intervals specially eon-
sjiieuous 2
2. Klytra with an interval (the 19th, near
humeri) more conspieuons than others at
base; see also Description (p. 220) Iniincidlis
— Elytra with no single interval more eoii-
spieuous than others at base 3
.3. Prothorax subeordate, with sides not oi'
weakh' sinuate; pronolum more evcuK pmie-
tate, the punctures rather coarse and of c.
uniform size (p. 221) . aiistidlis
- I'rothora.x strongly cordate, with sides stronglj'
sinuate; pronotum less evenly punctate, with
coarsi' and fine punctures mixed (p. 221)
cordens
Planetes secernendus OberthUr
Oberthur 1883, Notes Leyden Mus. 5, p. 217.
Description ( for recognition only ) . Form
small; sparsely inconspicuously pubescent;
piceous, not spotted, appendages brownish
testaceous; prothorax cordate or subeordate,
pronotum unevenly punctate, the punctures
varying in size and usually sparser near
middle of pronotum; elytra each with more
than 20 fine costae, the costae subequal
except 1st, 4th, 7th, etc. usualh' sliii^lithj
wider or more prominent toward base and
apex, but 1 9th costa not specially con-
spicuous at base; length r. 6.0-7.5 mm.
Types. From Sumatra; in Oberthiir
Coll., Paris Mus. (not seen).
Occurrence in New Guinea. Sixty-three
specimens from localities (including Dobo-
dura) scattered over most of the length of
New Guinea; at low altitudes only, none
above 500 in.
Notes. P. secernendus is now known
from the Malay Pen. (British Mus.),
Sumatra, Java (British Mus.), Borneo.
Leyte and Luzon in the Philippines
(M.C.Z.), New Guinea, and New Britain
(M.C.Z. ). Geographic \;uiation probably
occurs but is confused 1)\ iiidi\idual \aria-
tion especialK (in New (Guinea) in form
and pimctation ol prothorax. See also Addi-
tional material and Notes under /'. humer-
al is. below.
Plonefes humeralis n. sp.
nescri))lion. With characters ol genus;
ionn as in preceding species (seeenioidus),
reddish biown, sometimes darkcM', append-
ages slightK i^aler; head and pronotum
shining betwcHMi piuictures, eKlia duller.
lle(ul 0.77 and 0.7(S w idth prothorax, \\ eakly
impressed across base; I rout comex, slightly
impressed anleriorly, slightK irregularly
The Carabid Beetles of New Guinea • Darlington
221
finely punctate. Protlwrax narrowly sub-
cordate; width length 1.25 and 1.26; base
apex 1.06 and 1.05; sides broadly arcuate
anteriorly, moderately sinuate posteriorly,
with moderate margins, each with usual 2
setae; disc slightly convex, with middle line
well impressed but lateral longitudinal im-
pressions vague; baso-lateral impressions
moderate, roughened; surface of disc mod-
erately punctate with punctures of mixed
sizes, more closely punctate across base and
apex. Elytra subparallel; width elytra pro-
thorax 1.35 and 1.37; apices obliciuely trun-
cate, outer angles broadly rounded, sutural
angles scarcely blunted; each elytron with
more than 20 fine costae, the 1st, 4th, 7th,
etc., slightly more prominent than others
and the 19th specially prominent (but still
fine) at base. Secomkinj sexual characters:
5 front tarsi slightly dilated, 3 segments 2-
seriately squamulose; 1 principal seta each
side last ventral segment in both sexes.
Measurements (of types): length 7.3-8.3;
width 2.5-2.9 mm.
Types. Holotype c^ (Bishop Mus.) and
1 £ paratype (M.C.Z., Type No. 31,515)
from Eliptamin Vy., N-E. N. G., 1200-1350
m, July 16-31, 1959 (W. W. Brandt); 1
6 paratype, Torricelli Mts., Mokai Village,
N-E. N. G., 750 m, Jan. 1-23, 1959 (W. W.
Brandt, Bishop Mus.); 1 $ paratype, Mt.
Dayman, Maneau Rge., Papua, 700 m, "N.
Slope No. 6," July 13-20, 1953 (Geoffrey
M. Tate, A.M.N.H.).
Additiorml material. N-E. N. G.: 3,
Krisa, Vanimo, Apr. 1939 (Cheesman).
West N. G.: 1, Dojo, 2nd Strip, Res. Hol-
landia, Apr. 15, 1957 (R. T. Simon Thomas,
Louwerens Coll.).
Measured specimens. The 6 holotype and
i paratype from Eliptamin Vy.
Notes. The diagnostic character of this
species is the relative conspicuousness of
one costa (the 19th) at base of each elytron.
Form, color, punctation, and size are also
characteristic of the types. However, the
individuals listed under Additional material
are darker and much smaller than the types,
c. 6.5 mm long. They have the 19th costae
relatively conspicuous, as in the types, but
otherwise are more like secernendus. More
material from more localities is needed to
show wliether these specimens are referable
to humeralis or to secernendus, or whether
they represent a separate species. One
possibility is that humeralis occurs prin-
cipally at higher altitudes than secernendus
and that intermediates occur where their
ranges overlap.
Planeies ausfralis (Macleay)
Macleay 1871, Trans. Ent. Soc. New South Wales
2, p. 82 (PoUsticiis).
Description. None required here; note
size, prothorax with sides weakly or not
sinuate; pronotum c. evenly rather coarsely
punctate; length (in New Guinea) 7.7-9.5
mm.
Type(s). From Rockhampton, Queens-
land, Australia; presumably in Macleay
Mus., Sydney (not seen).
Occurrence in New Guinea. Papua: 1,
L. Daviumbu, Fly R., Sept. 11-20, 1936
(Archbold Exp., A.M.N.H.). West N. G.:
1, Kota Nika, Res. Hollandia, Jan. 25, 1956
(R. T. Simon Thomas, Louwerens Coll.),
in light trap; 1, Hoi Maffin, near Sarmi, July
18, 1959 (T. C. Maa, Bishop Mus.).
Notes. The New Guinean specimens
agree reasonably well with specimens from
Cairns, North Queensland, identified as
australis from description.
Planetes cordens n. sp.
Description. With characters of genus;
form as in Figure 134, depressed, with
wide-cordate prothorax; reddish piceous,
appendages paler; surface inconspicuously
pubescent (as usual); head and pronotum
shining between punctures, elytra dull.
Head 0.76 and 0.79 width prothorax, ir-
regularly impressed across base; front con-
vex except irregularly impressed anteriorly,
with a little irregular fine punctation. Pro-
thorax: width length 1.42 and 1.44; base
apex 0.97 and 0.96; sides broadly rounded
anteriorly, strongly sinuate posteriorly, mod-
222 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
erately margined, with usual 2 setae; pos-
terior angles well defined, right or slightly
obtuse; pronotum weakly convex, with well
impressed middle line and less distinct lon-
gitudinal impressions each side nearer mar-
gin than middle; baso-lateral impressions
shallow, closely microreticulate; surface of
disc otherwise rather closely punctate with
mixture of moderate and minute punctures.
Eli/fra subparallel; width elytra prothorax
1.34 and 1.30; apices obliquely truncate
with outer angles broadly rounded and in-
ner angles scarcely blunted ( as usual ) ;
each elytron with more than 20 fine costae,
the 1st, 4th, 7th, etc. slightly more promi-
nent than others, but no costa specially con-
spicuous at base. Secondary sexual char-
acters as for humcralis (2nd species above).
Measurements: length c. 9.0-9.5; width
3.2-3.3 mm.
Types. Holotype $, (Hungarian National
Mus.) and 3 paratypes (2 in M.C.Z., Type
No. 31,516) from Madang ("Friedrich-
W'ilh.-hafen"), N-E. N. G., 1896 (Biro);
and additional paratypes as follows, all
from N-E. N. G. ( Biro ) : 1, Stephansort,
Astrolabe Bay, 1900; 1, Simbang, Huon
Gulf, 1899; 1, Erima, Astrolabe Bay, 1897.
Meastired specimens. The S holotype and
1 9 paratype from Stephansort.
Notes. Among New Guinean Planetes,
this should be immediately recognized by
rather large size and wide-cordate prothorax
with disc flatter than usual and punctate as
described. Why Biro should have found
this species at four localities while no one
else has found it anywhere is a mystery.
Perhaps he obtained his specimens in a
special habitat b\' sjx'cial collecting methods,
perhaps by sitting leaf-debris from the
ground in rain forest.
Tribe HELLUODINI
Csiki 1932, Coleop. Cat., Carabidae, Harpaliiiac 7,
p. 1571 (sec for synoiiyiiiy and additional iclcr-
cnces ) .
This is a small tribe, confined to thv
tropical Asiatic-Australian area. ()iil\- 3
genera are recognized, of which only 1 is
represented in New Guinea and ( northern )
Australia. However, Iloloponerus fiodef-
froyi (Fairmaire) (1881, Le Naturaliste 3,
p. 381; 1883, Ann. Soc. Ent. Belgium 27, p.
2) of New Britain, although considered a
lebiine by Fairmaire and listed as one in
the Coleopterorum Catalogus (Csiki 1932,
p. 1361 ), may belong in this tribe. I do not
know this insect, but the description is of a
large carabid (perhaps the largest member
of the family in New Britain), 28 mm long,
with long mandibles, prothorax expanded
at sides, elytra sinuate-truncate and not
spined, and head at base with a strong
spine on each side. This description sug-
gests a very large Poi:,onoglossus-\ike carabid
with genae, which are prominently angu-
late or tuberculate in some Pogonoglossus,
produced as spines.
Genus POGONOGLOSSUS Chaudoir
Chaudoir 1862, Bull. Soc. Nat. Moscow 35, Part
2, p. 304.
Csiki 1932, Coleop. Cat., Carabidae, Harpaliiiae 7,
p. 1571 (see for synonymy and additional refer-
ences ) .
Andrewes 1937, Bull. Soc. Ent. France for 1937, pp.
152// (with key to species of Ja\a and Sumatra).
Diaiinosis. Form including form of eyes
characteristic; upper surface at least partly
pubescent; antennae not geniculate, with
moderate 2nd segments; see description of
ligula and paraglossae, below.
Description (characters common to New
Guinean species). Form as in Figures 135-
140; variably pubescent. Head: eyes ±
abruptly prominent, genae rounded or an-
gulate-tubereulate behind eyes; 2 setae over
each eye; antennae not gtmiculate, 2nd seg-
ments moderate (not \('r\ short), segments
1-4 variably setulose, outer segments mor(>
densely pub(>seent; mandibles long, weakly
arcuate; neck deeply transxcrseK con-
stricted; front 2-impressed anterior!) ;
clypeus irregularly truncate, apparenth
2- or 4-setose anteriorU- (setae difficult
to distinguish Ironi other jMibescence );
labium \ ariable. 6-sc>tose; ineiituin with
The Carabid Beetles of New Guinea • Darlington 223
triangular tooth; ligiila short, rounded, with
c. 4 setae at apex and 2 more near middle
of length; paraglossae very slender, much
longer than and free from ligula; palpi
with apical segments narrowly subtruncate.
Prothorax cordate or subcordate; apex
slightly or moderately (not deeply, in New
Guinean species) emarginate, with anterior
angles usually rounded ( ± pointed in some
g,lobncoUis) and not or not much advanced
beyond arc of emargination; base subtrun-
cate or emarginate at middle, ± oblique at
sides; posterior angles or sides of prothorax
just before angles usually minutely emar-
ginate; margins moderate or wide, reflexed,
each with seta at base and before middle;
disc usually only weakly convex, with mid-
dle line and transverse impressions distinct,
baso-lateral impressions present Init not
sharply defined. Elytra quadrate; margins
usually faintly subserrate; apices obliquely
emarginate-truncate with membranous mar-
gins, with outer angles usually rounded
(obtuse in popiio), inner angles c. acute or
blunted, not armed; striae entire ( ± obso-
lete in unicolor and globricoUis); intervals
variably punctate, 3rd with up to 3 or 4
special dorsal punctures (often not distin-
guishable especially in species with exten-
sive general punctation). Inner uiniis full.
Lower siu-jace variably punctate-pubescent.
Legs moderately slender; tarsi pubescent
above, not sulcate above; 4th hind-tarsal
segments shallowly emarginate; 5th seg-
ments setulose above and below; claws
simple. Secondary sexual characters: S
front tarsi not or not much widened, 3 seg-
ments narrowly 2-seriately squamulose; S
usuallv with 2 or 3, 9 3 or 4 setae each
side last ventral segment, but these setae
and their punctures sometimes difficult to
distinguish.
Type species. P. validicornis Chaudoir,
of Java.
Generic distribution. SE. Asia to north-
ern Australia.
Notes. Species of this genus are probably
moderatelv numerous and diverse from the
SE. corner of Asia to New Guinea (fewer
in northern Australia), but individuals are
rarely collected. Of 9 Javan and Sumatran
species, Andrewes ( 1937 ) saw only single
specimens of 6; and of 9 (or 10, with
unicolor (Macleay) ) New Guinean species,
I have seen a satisfactory series of only 1.
All the New Guinean species appear to be
endemic. I compared some of them with
the Andrewes Collection in 1948 ( see Notes
under several species, below); none fits
the description of P. horni Sloane ( 1907,
Deutsche Ent. Zeitschrift for 1907, p. 184)
of New Britain; and the 2 Australian species
that I have seen are different from any New
Guinean species.
Key to Species of Pogonoglossus of
New Guinea
1. Elytra] striae distinct, impressed 2
- Elytral striae faint or absent 9
2. Genae rounded or irregular behind eyes but
not conspicuously angulate or tuberculate
(see Description of taylori) 3
- Genae conspicuously angulate or tuberculate . 7
3. Entire upper surface including much of
head rather closely punctate or (on elytra)
roughened 4
- Part or all of upper surface sparsely punctate
or impunctate 6
4. Elytra with outer-apical angles obtuse but
distinct; size medium (length 9.0-11.5 mm);
(found at low altitudes) (p. 224) painia
- Elytra witli outer-apical angles rounded;
size either larger or smaller; (often at higher
altitudes ) 5
5. Larger, length 12.3-13.0 mm (see also De-
scription) (p. 224) taylori
- Smaller, length 7.6-8.7 mm (see also De-
scription) (p. 225) ininor
6. Prothorax less wide (width/length 1.49),
with moderate margins (p. 225) major
- Prothorax very wide (width/length 1.88 and
1.97), with very wide margins (p. 225) latior
7. Sides of prothorax oblique but scarcely sinu-
ate posteriorly; length c. 13 mm (p. 226) -
obliquus
- Sides of prothorax sinuate posteriorly;
smaller °
8. Length 9.6-11.0 mm (p. 226) grossuliis
- Length 7.0-9.0 mm (p. 227) parvus
9. Pronotum densely minutely punctate and
pubescent (p. 227) unicolor
- Pronotum virtually impunctate and glabrous
(p. 227) glahricollis
224 BuUetin Museum of Comparative Zoology, Vol. 137, No. 1
Pogonoglossus papua n. sp.
Description. With characters of genus;
form as in Figure 135; brownish black, ap-
pendages dark; entire upper surface mod-
erately pubescent, punctate, with reticulate
microsculpture indistinct or (on elytra) ir-
regular. Head 0.84 and O.Sl width pro-
thorax; genae rounded, not strongly angu-
late; front moderately punctate, shining be-
tween punctures. Prothorax strongly cor-
date; width/length 1.45 and 1.48; base/apex
1.14 and 1.19; base broadly slightly emar-
ginate, slightly oblique at sides; sides
strongly sinuate well before base; basal
angles shaiply formed, c. right; margins
rather wide, moderately reflexed; disc
weakly convex, surface moderately closely
punctate-pubescent, less shining than head
but more shining than elytra. Elytra:
width elytra prothorax 1.46 and 1.46; outer-
apical angles obtuse but more distinct than
usual in genus, striae moderately impressed,
indistinctly punctulate; intervals slightly
convex, closely punctate-pubescent. Sec-
ondary sexual characters as for genus; S
with 2, 9 3 special seta-bearing punctures
each side last ventral segment. Mca.siire-
ments: length c. 9.0-11.5; width 3.4-4.1
mm.
Types. Holotype $ (M.C.Z., Type No.
31,517) and 10 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. Papua: 1,
without precise locality (Hungarian Na-
tional Mus.). N-E. N. G.r 1, Lae, sea level,
July 24, 1955 (Gressitt), in light trap; 1,
Busu H., E. of Lae, 100 m, Sept. 13, 1955
(Gressitt); 7, Aitape, Aug. 1944 (DarHng-
ton). WestN. G.: 1, Hollandia, July-Sept.
1944 (13arlingt(m); 2, same locality. May
1945 (II. Iloogstraal, M.C.Z.); 1, same lo-
cality, June 1945 ( B. Malkin, U.S.N.M.); 3,
same locality, dates in Nov., Dec, Jan.
1944-1945 (W. T. Nailon, Fenton Coll.);
1, Sentani, 90+ m, June 22, 1959 (Gressitt
and T. G. Maa, Bishop Mus.), in light trap;
1, Hoi Maffin, near Sarmi, July 18, 1959 (T.
G. Maa, Bishop Mus.); 1, Neth. N. G. with-
out further locality, Oct. 10, 1944 (T.
Aarons, Gal. Acad.). Also 1, "Sinimi" { =
Senimi R., Papua?), "Vr, 1943" (T. Niimura,
Ueno Goll.).
Measured specimens. The S holotype and
1 9 paratype from Dobodura.
Notes. In Andrewes' key ( 1937, see refer-
ence under genus), this species runs to
latus Andrewes of Sumatra but has the pro-
thorax probably narrower at base and less
emarginate at apex and the pronotum cer-
tainly more closely punctate. Of Australian
species, papua is closest to porosus Sloane
( I have specimens, identified from descrip-
tion, from Rocky R., mid-peninsular Gape
York ) but has a more strongly cordate pro-
thorax and better defined outer-apical
elytral angles.
Since papua is the common Poii,ono'jJos-
sus in New Guinea, I shall take it as a
standard for comparison of sexeral of the
following species.
Most of my Dobodiua specimens were
taken in piles of dead leaves on the ground
in rain forest.
Pogonoglossus taylori n. sp.
Description. With characters of genus;
form and characters c. as preceding species
{papua) except as follows. Head 0.15 and
0.78 width prothorax; genae more promi-
nent than in papua, nearly wide as eyes,
minutely tuberculate and e. subangulate be-
hind eyes. Prothorax: width length 1.44
and 1.58; base/apex 1.18 and 1.18; apex
slightly more emarginate than in ))apua
and sides slightly more broadl\ and e\enl>
reflexed. Elytra: width elytra prothorax
1.43 and 1.43; outcM-apical angles more
rounded than in papua. Measurenwnis:
length 12.3-13.0; 4.3-4.9 mm.
Types. Holotype S (M.G.Z., l\pe No.
31,518) from Aiyura, N-E. N. G.. 'l900 m
July 1962 (R. W. Taylor, #2147), in rain
forest; 1 9 paratvpe, Eliptanu'ii \\'., N-E.
N. G., 1665-2530 m, June 23-30. 1959 (W.
W. Brandt, Bishop Mus.); 1 s paratvpe,
Okapa, N-E. N. G., Aug. 29, 1965 (R.
Hornabrook).
The Carabid Beetles of New Guinea • Darlington
225
Measured specimens. The i holotype and
9 paratype from Eliptamin Vy.
Notes. This is apparently a mountain-
Hving species probably related to the low-
land papiia but differing from it as indi-
cated in the Description above.
Pogonoglossus minor n. sp.
Description. With characters of genus
(except i unknown); form c. as in papua;
characters c. as in papna except as follows.
Color browner (less black), surface slightly
more shining. Head 0.88 and 0.86 width
prothorax; eyes slightly smaller and genae
more evenly rounded than in papiia. Pro-
thorax: width length 1.47 and 1.49; base
apex 1.08 and 1.07. Elytra: width elytra
prothorax 1.50 and 1.51; outer-apical angles
more rounded than in papiia; intervals less
roughened. Measurements: length 7.6-8.7;
width 3.0-3.3 mm.
Types. Holotype 9 (M.C.Z., Type No.
31,519) from lower Busu R., Huon Pen.,
N-E, N. G., May 12, 1955 (E. O. Wilson),
in lowland rain forest; 1 9 paratype, W'au,
Morobe Dist., N-E. N. G., 1200 m, June 22,
1961 (Sedlaceks); 1 9 paratype, Hollandia,
West N. G., Jan. 20, 1945 (W. T. Nailon,
Fenton Coll.); 1 9 paratype, Njau-limon,
S. of Mt. Bougainville, West N. G., 300 ft.,
Feb. 1936 (Cheesman).
Measured specimens. The 9 holotype and
9 paratype from Njau-limon.
Notes. P. minor differs from papua as
indicated in the preceding Description.
The 2 species are sympatric but minor is
apparently the less widely distributed,
being known only from a comparatively
small part of east-central New Guinea.
P. minor is similar also to porosus Sloane
of North Queensland, Australia, but has the
head more punctate and the prothorax more
strongly cordate.
Pogonoglossus major n. sp.
Description. With characters of genus;
form as in Figure 136; irregular brownish
piceous, appendages dark brown; rather
shining, reticulate microsculpture absent or
indistinct on head and pronotum, light,
irregular, moderately transverse on elytra;
surface punctate as described below. Head
0.82 width prothorax; genae prominently
rounded but not angulate; front sparsely
punctate-pubescent. Prothorax weakly cor-
date; width/length 1.49; base/apex 1.05;
sides broadly sinuate before obtuse except
minutely subdenticulate posterior angles;
surface irregularly rather sparsely punctate-
pubescent. Elytra: width elytra/prothorax
1.44; outer-apical angles broadly rounded,
sutural angles blunted; striae deep, entire,
finely punctulate; intervals convex, sparsely
punctate, 3rd with apparently 3 or 4 special
dorsal punctures difficult to distinguish
from other punctures. Secondary sexual
characters: S front tarsi as genus; 6 with
apparently 3 principal setae on left, 4 on
right side last ventral segment; 9 unknown.
Measurements: length 17.5; width 6.0 mm.
Type. Holotype S (M.C.Z., Type No.
31,520) from vie. Nadzab, N-E. N. G., July
1944 (Darlington); the type is uniciue.
Notes. This is the largest New Guinean
Pogonoglossus. It is about the size of P.
Jwrni Sloane (Deutsche Ent. Zeitschrift for
1907, p. 184) of New Britain but has the
prothorax more narrowed behind with more
obtuse posterior angles, the outer elytral
striae not fainter, and the elytral intervals
sparsely rather than closely setose-punctate.
Pogonoglossus lafior n. sp.
Description. With characters of genus;
form as in Figure 137, very broad; brownish
black, appendages dark; moderately shin-
ing, reticulate microsculpture indistinct on
head and pronotum, light, fine, rather
strongly transverse on elytra; punctation as
described below. Head 0.74 and 0.67 width
prothorax; genae oblique for most of length,
slightly rounded or very obtusely subangu-
late behind eyes; front c. impunctate ex-
cept sparsely punctate laterally and pos-
teriorly. Prothorax very wide, cordate;
width length 1.88 and 1.97; base/apex 1.24
226 Btilletin Museum of Comparative Zoology, Vol. 137, No. 1
and 1.12; sides strongly but variably sinuate
well before c. right or obtuse posterior
angles; margins widely reflexed; disc more
convex than usual, sparsely inconspicuously
punctate-pubescent. Elytra: width elytra
prothorax 1.45 and 1.30; outer-apical angles
rounded, sutural angles blunted; striae
entire, well impressed, slightly irregular
but scarcely punctulate; intervals convex,
sparsely inconspicuously punctulate, 3rd
with 3 or 4 dorsal punctures difficult to
distinguish. Sccundary sexual characters: 6
front tarsi as for genus; 6 with 3, 9 4 setae
each side last ventral segment. Measure-
ments: length 14.5-15.5; width 5.7-6.0 mm.
Types. Holotype c5 ( Leiden Mus. ) and
1 9'paratype (M.C.Z., Type No. 31,521)
from Lower Mist Camp, Snow Mts., West
N. G., 1550 m, Jan. 31, 1939 (Toxopeus).
Notes. See Key to Species for distinguish-
ing characters of this well defined species.
The 9 has a wider prothorax with more
obtuse angles than the S , but this is prob-
ably individual rather than sexual variation.
I have no doubt the 2 specimens are con-
specific.
Pogonoglossus obliquus n. sp.
Description. With characters of genus;
form as in Figure 138; black, appendages
dark; shining, reticulate microsculpture
absent or indistinct even on elytra; puncta-
tion as described l)elow. Head 0.77 width
prothorax; genae prominently angulate-
tuberculate behind eyes; front almost
smooth, very sparsely punctulate-pubescent.
Prothorax very wide; width/length 2.00;
base/apex 1.22; sides oblicjue and converg-
ing and scarcely sinuate before obtuse pos-
terior angles; margins widely reflexed; disc
moderately convex, sparsely punctate-jm-
bescent, more closely so across base and
apex. Ehjtra: width not measured (speci-
men broken); humeral margins wider ihan
usual; outer-apical angles roimded, sutural
angles acute, scarcely blunted; striae entire,
impressed, irregular but seareel\- punctu-
late; interxals con\-ex, finely s])arsel\- punc-
tulate, 3rd with c. 3 special dorsal punc-
tures difficult to distinguish. Secondary
sexual cJiaracters: 6 front tarsi as for
genus; i with 3 setae each side last
ventral segment; 9 unknown. Measure-
ments: length c. 13 mm; width not mea-
sured.
Ty])e. Holotvpe 6 (Bishop Mus.) from
Eliptamin Vy.,'N-E. N. G., 1665-2530 m,
June 23-30, 1959 (W. W. Brandt); the type
is unique.
Notes. The single specimen was received
in bad condition and remounted in pieces
on a card, but it shows the essential char-
acters of this very distinct species. See Key
to Species for its differential characters.
Pogonoglossus grossulus n. sp.
Description. With characters of genus;
form average; black or brownish black, ap-
pendages dark; shining, reticulate micro-
sculpture indistinct even on elytra. Head
0.80 and 0.79 width prothorax; genae promi-
nently angulate-tuberculate behind eyes;
front virtually smooth at middle, very
sparsely punctulate-setose at sides. Pro-
thorax wide-cordate; width length 1.95 and
1.94; base/apex 1.23 and 1.19; sides broadly
sinuate before c. right or obtuse posterior
angles; margins wide, widely reflexed; disc
weakly convex, very sparsely punctulate-
pubescent. Eh/tra: width elytra prothorax
1.27 and 1.36; outer-apical angles rounded,
sutural angles acute (except for membranous
margins); striac> deep, scarcely punctulate;
intervals convex, very sparsely punctulate,
3rd with up to 3 special dorsal punctures
difficult to distinguish. Secondary sexual
characters as for genus; S with 2, 9 with 3
setae each side last ventral segment. Mea-
surcmoils: length 9.6-11.0; width 3.7-4.1
mm.
Types. Holotype 9 (M.C.Z., Type No.
31,522) Irom \ic. ZcMigarn, \'\-. of Kua H.,
Mongi Watershed, Ihion Pen., I\-K. N. (i..
800 m, Apr. 14, 1955 (E. O. Wilson); 1
9 paratype, Lae, N-E. N. (i.. JuK 1941
(F. E. Skinner, Purdue V. Coll., borrowed
fr. Bisho]-) Mus.); 1 ^ paratope. Kokoda.
The Carabid Beetles of New Guinea • Daiiington 227
Papua, 1200 ft. (366 m), Sept. 1933 are distinet in both specimens and may
(Cheesman). prove to be characteristic of the species,
Measured specimens. The ^ paratype and although similar marks are indicated in
9 holotype, in this order. some individuals of some other species.
Notes: Although the 3 individuals listed
above vary somewhat, they agree in form Pogonoglossus unicolor (Macleay)
of genae, wide-cordate prothorax, shining Macleay 1886, Proc. Linnean Soc. New South
sparsely punctate surface, and moderate Wales (2) 1, p. 137 {Planetes).
size. I think they probably represent a ^'"''"'^ ^^^'^^ Deutsche Ent. Zeitschrift for 1907,
single, variable species.
Description ( significant details only, from
Pogonoglossus parvus n.sp. Macleay's description). Color brownish
Description. With characters of genus; hlack, legs dark; head shining, pronotum
form as in Figure 139; brownish black, head and elytra dull and densely minutely punc-
with 2 narrow oblic^ue red marks posteriorly tate; prothorax a little wider than long, with
(see following Notes), appendages dark titles narrowed to posterior angles which
brown; moderately shining, reticulate micro- ^re "rather obtusely rectangular"; elytra
sculpture indistinct on head, irregular or "with 8 or 9 almost invisible striae"; length
transverse and light on pronotum and elytra, c. 10 mm.
Head 0.85 and 0.85 width prothorax; genae Type. From Fly R. (probably Papua);
angularly prominent behind eyes; much of should be in Macleay Mus., Sydney (not
front virtually impunctate. Prothorax cor- seen).
date; width/length 1.48 and 1.56; base apex Notes. Sloane (1907) adds nothing to
1.21 and 1.23; sides broadly sinuate before Macleay's description of unicolor except
obtuse or nearly right posterior angles; ^^'^^ the insect is a Pogonogjo.ssus. The
margins rather narrow and not strongly re- ^^^Y lightly striate elytra distinguish it from
flexed; disc moderately convex, finely ^H known New Guinean species of this
punctate-pubescent ( pubescence rubbed off genus except glabricoUis Van Emden ( be-
in part in holotype ) . Elytra: width elytra low), from which it differs in having the
prothorax 1.34 and 1.35; outer-apical angles pionotum densely punctate and pubescent
narrowly rounded, sutural angles acute or rather than smooth and virtually glabrous
slightly blunted; striae well impressed, ir- '^■'^ i" glabricoUis.
regular but scarcely punctulate; intervals „ ; ; , • //■ v, .- ■
r.r^r.,.^v. c. 1 1^ 1 4. 1 i. o 1 Pogonoglossus glabncollis Van Emden
convex, sparsely punctulate-pubescent, 3rd ^ ^ »
^^'ith special dorsal punctures not surely ^''" ^'"''™ ^•^•^"' ^^^""^^■•" E"^- ^^^^""'^ 9^' P- ^^■
distinguishable. Secofu/f/rj/ .sexjva/ c/i<3/rtcfers Description. With characters of genus;
as for genus; c^ with 2 or 3 (unsymmetric), form as in Figure 140 (but somewhat vari-
9 3 setae each side last ventral segment, able); irregular dark brown or brownish
Measurements: length 7.0-9.0; width 2.5- black, head with 2 reddish marks posteri-
3.2 mm (the 6 is the larger). orly, appendages brown; moderately shining,
Types. Holotype $ (M.C.Z., Type No. elytra duller, reticulate microsculpture in-
31,523) from vie. Hollandia, West N. G., distinct on head and pronotum, light and
July-Sept. 1944 (Darlington); and 1 9 irregular on elytra. Head 0.85 and 0.83
paratype, Dobodura, Papua, Mar.-July 1944 width prothorax; genae rounded; front
(Darlington). almost impunctate. Prothorax cordate, vari-
Notes. The small size, angulate genae, able; width/length 1.39 and 1.60 {sic);
and form and microsculpture distinguish base/apex 0.98 and 1.07; sides broadly sinu-
this species. The red marks on the head ate before obtuse or c. right posterior
228 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
angles; margins rather narrow l:)iit variable;
anterior angles rounded or bluntly pointed;
disc almost flat, scarcely punctulate. Elytra:
width elytra/prothorax 1.35 and 1.37; outer-
apical angles broadly rounded, sutural
angles acute or blunted; striae faintly in-
dicated or virtually obsolete; surface closely
punctulate; up to 3 apparent dorsal punc-
tures sometimes visible on position of 3rd
intervals. Secondary sexual characters un-
determined ( i unknown); 9 with 3 or 4
(sometimes unsymmetric) setae each side
last ventral segment. Measurements: length
12.5-16.0; width 4.2-5.3 mm.
Type. A 9 from N-E. N. G.; in Van
Emden Coll., British Mus. (seen).
Occurrence in New Guinea. Papua: 1
9, Kokoda-Pitoki, 400 m. Mar. 23, 1956
(Gressitt); 1 9, Fiume Purare, Jan. 1894
(Loria, borrowed from Straneo). N-E.
N. G.: the holotype; 1 9, Motae, Kuku
Kuku, E. Highlands, 6000 ft. (c. 1830 m),
"1/3/64" (R. Hornabrook). West N. G.: 1
9 , Geelvink Bay, 1878 ( Raffray and Main-
dron, Paris Mus.).
Measured specimens. The 9 9 from
Kokoda-Pitoki and Motae.
Notes. Although the 4 individuals listed
above vary in several characters ( e.g. form
of prothorax, degree of obliteration of
elytra! striae), the variations are not obvi-
ously concordant, and I think only one very
distinct species is involved. It is uniquely
characterized by form, elytral striae faint or
obsolete, and combination of virtually im-
punctate jironotum and densely punctulate
elytra.
Tribe HELLUONINI
Sloanc 191 I, I'loc. Liiiiican Soc. New Sontli Wales
39, p. 568.
Csiki 1932, Colcop. Cat., Carabidac, liarpalinac 7,
p. 1572 (see for synonymy and additional rcfcr-
fnccs).
Jeanne! 1949, Colcop. Carahiciucs dc la Hejjion
MaljJiaclie, Part 3, p. 1041.
Jrdlicka 1963, Knt. Abliandlnn«en 2S, p. 167.
llclliioiiinac Basilfwsky 1953, Exploration Pare
National I'Upemba, Fasc. 10. p. 219.
This is still another small but widely
distributed tribe. The members of it are
medium-sized or large carabids, usually of
characteristic form, usually with sparse or
short pubescence, and usually with mouth-
parts including the labrum strikingly modi-
fied. Three genera are confined to the
Americas; 6, to Africa and/or the Oriental
Region (except that a species of Crea^.ris
extends to Australia); 13, to the Australian
Region. (A supposed helluonine on Ne\v-
Caledonia has been shown not to be one
by Britton, 1937, Ent. Monthly Magazine
73, p. 127. ) The Australian genera form a
distinct group of the tribe, characterized by
Sloane (1914, p. 570). Five genera (1 of
them new) and 8 species of Australian-
group Helluonini occur in New Guinea,
where the only other member of the tribe
is CreaiS,ris lahrosa, \\'hich ranges frotn
Ceylon and India to Australia.
In spite of Sloane's ( 1914 ) careful stud\
of the Australian genera, I have had trouble
with the generic classification of the New
Guinean forms. This is partK' because m\'
material is inadequate: 2 obviously distinct
new species are represented by unique fe-
males which I have assigned to Ilelhionidius
with some doubt, and I have been forced to
base an apparent new genus on a single
male. I myself found no Helluonini during
11 months in New Guinea and I can sa\
nothing about their habitats or habits there
except that all the New Guinean species
are winged and that some of them fl\' to
light. In Australia, different hc41uonines
lixc on the ground and on tree trunks, usu-
alK in open or openly-wooded places rather
than in rain forest.
K'i;v lo (Iknkha ok IIki.ia omm or Xi:w Cvinka
1. Front iemora not anuniate-protnlicrant he-
low: size smaller, lenutli r. 9 mm (p.
229 ) _ - - Creofiris
- Front femora tliickened and lilnntly angn-
late or protnl)eranl hi'low near base; size
larjier — - 2
2. Prothorax inoderateK' narrowed posteriorly,
witii si(.les moilerately siimate (p. 2'v3 ) _,
IIcIIikkIciiki
The Carabid Beetles of New Guinea • Darlington
229
- Piothoiax strongly narrowed posteriorly, with
sides strongly sinuate and base subpeduncu-
late 3
3. Ligula subtriangular, narrowed anteriorly,
with apex narrowly rounded ( p. 233 )
Helhiosoma
- Ligula very wide, with apex broadly rounded
or eniarginate 4
4. Labrum with 2 principal setae; elytra with
Sth intervals much wider tlian 7th and closely
punctate; length ( in New Ciuinea ) c. 30 mm
( p. 233 ) Gigadema
- Lal)rum with 4 or more principal setae; elytra
with 8th intervals not much wider than 7th
and less closely punctate; length c. 20 mm
or less 5
5. Tarsal segments unusually widened or par-
allel-sided, the 4th hind-tarsal segments eniar-
ginate for more than V2 the segments' length;
labrum usually with 4 principal setae (p.
229 ) Helluoniduis
- Tarsal segments not thus widened and not
parallel-sided, the 4th hind-tarsal segments
shallowly emarginate; laJMum with c. 10
principal setae (see also Description) (p.
232 ) HcUuopapud
Genus CREAGRIS Nietner
Nietncr 1857, J. Asiatic Soc. Bengal 26, p. 139.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1575 (see for synonymy and additional refer-
ences ) .
Diagnosis. See preceding Key to Genera.
Description. None required here.
Type species. Creagris lahrosa Nietner,
below.
Generic distribution. Six species in the
Oriental Region, 1 of them extending to
New Guinea and Queensland, Australia;
possibly an additional species in Queens-
land.
Notes. The listing of C. wiJsoni Castelnau
(the supposed endemic Queensland species)
also from Java by Csiki (p. 1576) is ap-
parently a compiler's error based on a mis-
reading of Sloane 1914 ( see reference under
following species ) .
Creagris labrosa Nietner
Nietner 1857, J. Asiatic Soc. Bengal 26, p. 139.
Sloane 1914, Proc. Linnean Soc. New South Wales
39, p. 570.
1920, Proc. Linnean Soc. New South Wales
45, p. 322.
Csiki 1932, Coleop. Clat., Carabidae, Harpalinae 7,
p. 1575 (see for synonymy and additional refer-
ences ) .
Description. None required here; note size
small; color dark brown; labrum expanded,
r. circular, shallowly channeled each side;
mentum with lateral lobes and median
tooth all produced as long very slender
processes; front femora not angulate below;
length c. 9 mm.
Types. From Ceylon; in Berlin U. Zool.
Mus. and Stettin Mus. (not seen).
Occurrence in New Guinea. Papua: 2,
Mt. Lamington, 1300-1500 ft. (c. 400-460
m) (C. T. McNamara, S. Australian Mus.).
Notes. C. lahrosa is now known from
Ceylon, India, Burma, etc., Java, New
Guinea, and Queensland, Australia (a
specimen from Mackay, recorded by
Sloane, 1914).
Genus HELLUONIDIUS Chaudoir
Chaudoir 1872, Revue et Magasin Zool. (2) 23,
p. 216.
Sloane 1914, Proc. Linnean Soc. New South Wales
39, pp. 571, 582.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1580 (see for synonymy and additional refer-
ences ) .
Diagnosis. Form c. as in Figure 142;
pubescent (as usual in tribe); genae vari-
able; labrum variable, produced or angu-
late at apex, usually with 4 principal setae;
mentum toothed; ligula rounded; prothorax
strongly constricted at base; elytra with Sth
intervals not much wider than 7th, irregu-
larly punctate; 4th hind-tarsal segments ±
wide, deeply emarginate; see also preceding
Key to Genera of UeUuonini of New Guinea.
Description. None attempted here; ma-
terial inadequate.
Type species. Aenigma cyanipcnne Hope,
of Australia.
Generic distribution. Eastern and north-
ern Australia, New Guinea.
Notes. Of the 4 New Guinean species
now placed in this genus, only chrysocomes
Maindron is a typical Helluonidius. The
other 3 species, 2 of them based on unique
230 Bulletin Museum of Coniptudfive Zoology, Vol. 137, No. 1
females (sex determined by dissection),
differ in form of labrum and differ among
themselves in form of genae, form of tarsi,
and in other ways. They are obviously
distinct species, but males are needed to
determine generic assignments.
Key to Species of Helluonidius of
New Guinea
1. Lalnuni loiijier than wide, narrowly pro-
duced at apex (p. 230) chrysocoitics
- Labrum wider than long, not narrowly pro-
duced at apex 2
2. Labrum witli median setae close to margin;
cKtra microreticulate (p. 230) laevifrons
- I^abrum with median setae c. V:\ lalirum's
length behind margin; elytra not micro-
reticulate 3
3. Genae abruptly truncate, c. straight from
posterior edges of eyes to neck; tarsi \'er>'
wide (Fig. 184) (p. 231) latipes
- Genae moderately convex; tarsi less wide (p.
231) politus
Helluonidius chrysocomes Maindron
Mainchon 1908, Ko\a (Guinea 5, Li\raisf)n 2, p.
299.
Description. None required here; form
as in Figure 142; form of labrum ( longer
than u'ide, and narrowK' produced at apex )
unique in tribe Ilelluonini in New Guinea;
genae prominent; antenna and hind tarsus,
Figure 182; color black or dark brown; sur-
face without reticulate microsculpturc;
lengtli c. 16.()-1(S.5 mm.
Type. From "Sentani" (near llollandia ),
West N. G.; probably in Paris Mus. (not
seen ) .
Occurrence in Neic Cuineo. Twenty-two
speeimens from 11 localities scattered oxer
most ol the length of New (Guinea, from
I^)rt Moresby and the Fly R. to the N'ogel-
kop; at low iiltitudes, none labeled higluM"
than SOO ni ( Araucaria Camp).
Notes. This is a t\'pical IJelluoniditis,
very close to ;ind perhajis the same as one
of the Australian species, which are all \'(m\'
similar and which n(H>d revision.
Most spec iiiK'iis ol elinjsocomes haxc the
surface especially ol the elytra w ith a jiearly
luster which is apparently due to a surface
film of some sort, but fi individuals w ideU'
scattered within the range of the species are
shining black without luster.
Helluonidius laevifrons n. sp.
Description. With characters of genus;
form as in Figure 143; antenna and hind
tarsus. Figure 183; black, appendages dark;
head and pronotum shining (but punctate
as described below), elytra duller, with c.
isodiametric microsculpturc. Head 0.84
width prothorax; genae oblique, not at all
prominent; chpeus subtrimcate, 1 -setose
each side; labrum wider than long, obtuseK'
subangulate, sinuate each side of angula-
tion, 4-setose with inner setae very close to
anterior margin (see Notes, below); front
deeply impressed each side, the impressions
punctate but front otherwise broadh' im-
punctate; neck moderately impressed and
punctate; mentum with large triangular
tooth; ligula wide, irregularly rounded, de-
pressed at middle, with 3 pairs seta-bearing
punctures one behind the other; inner lobe
of maxillae with inner edge irregular near
middle and with strong hook at c. right
angles from inner-apical angle; palj^i rather
short and thick. Protliora.x: width length
1.19; base/apex 1.02; base head 0.72; mar-
gins narrow, scarcely interrupted; disc ir-
regularly convex, irregularly coarsely punc-
tate with punctures tcmding to form longi-
tudinal rows between n;urow smooth sp;ices
at middle. Elyfrd: width ebtra prothorax
1.22; striae well impressed, punctulate; in-
tervals slight 1\ convex, most int(M\als
slightly irrt\gularly 2-seriately punctate,
si)ccial dorsal punctures of 3rd not sureK
distinguish;ibl(\ 8th slightK' wider than 7th
and irregularly (not 2-seriately ) punctukite.
Lciis: tarsal segments more nearl\- j^arallel
than usual (Fig. 183); 4th hind-tarsal seg-
ments emarginate for slightK more* than ^-i
length. Measurcnicnls: length 18.0; width
6.5 mm.
Type. Il()l()t\pe V (Bishop Mus.) Irom
Torrieelli Mts.,'M()kai Village, N-E. N. G.,
750 m. j;ui. 1-23, 1959 (W. \\ . Brandt);
the type is unique.
Notes. The lionl maiuiii ol ihc ];i1)rum
The Carabid Beetles of New Guinea • Dadingion
231
is slightly sinuate on each side, with the
inner seta-bearing puncture near margin at
the sinuation. This form of lahrum is inter-
mediate between that of //. cJirysocomcs
and those of the following two species. But
in some other ways ( form of genae and
especially form of tarsi ) the present species
is strongly characterized, not intermediate.
Helluonidius lafipes n. sp.
Description. With characters of genus;
form as in Figure 144; antenna and hind
tarsus. Figure 184; dark brown, appendages
dark; shining, without reticulate microsculp-
ture, but punctate as described below.
Head 0.82 width prothorax; genae oblique,
not at all prominent; clypeus subtruncate,
2-setose each side; labrum wider than long,
bluntly obtusely angulate, not sinuate at
sides of angulation, 4-setose with inner setae
almost % length of labrum behind anterior
margin; front irregularly convex, deeply im-
pressed each side anteriorly, punctate at
sides and base but c. impunctate at middle;
mentum with strong bluntly triangular
tooth; ligula wide, rounded, scarcely im-
pressed at middle, probably 6-setose as in
laevifrons (above) but anterior setae
covered or broken; inner lobe of maxillae c.
as in laevifrom-, palpi stout. Frothorox:
width length 1.40; base apex 1.01; base '
head 0.72; margins narrow, much inter-
rupted anteriorly; disc irregularly convex,
irregularly punctate. Elytra: width elytra/
prothorax 1.52; striae impressed, scarcely
punctulate; intervals slightly convex, ir-
regularly 2-seriately punctate, 3rd with
special dorsal punctures not surely distin-
guishable, 8th not wider than 7th, irregu-
larly, rather sparsely in part 2-seriately
punctate. Le^s: tarsi exceptionally wide
(Fig. 184); 4th hind-tarsal segments much
wider than long, deeply and widely emar-
ginate; 5th segments wide and flattened.
Measurements: length 19.8; width 6.8 mm.
Type. Holotype 5 (Leiden Mus.) from
Rattan Camp, West N. G., 1200 m, Feb.-
Mar. 1939 (Toxopeus); the type is unique.
Notes. This species is assigned to
Helluonidius with doubt. The genae are
formed as in the preceding species (laevi-
frons) and the labrum is almost the same
in shape, but the 2 inner setae of the
labrum are much farther back and the tarsi
are strikingly different.
Helluonidius polifus n. sp.
Description. With characters of genus;
form as in Figiue 145; antenna and hind
tarsus. Figure 185; brownish black, append-
ages dark; shining, without reticulate micro-
sculpture but punctate as described below.
Head 0.84 width prothorax; genae moder-
ately convex, subprominent; clypeus broadly
slightly emarginate, 2-setose each side;
labrum wider than long, bluntly angulate,
with apparently 2 principal setae on right
and 4 on left, the inner seta on each side
almost V:'. length of labrum behind anterior
margin; front weakly convex, deeply im-
pressed each side anteriorly, punctate at
sides and across base and with a few widely
scattered punctures near middle; mentum
with strong blunt tooth; ligula wide,
rounded, scarcely impressed, 6-setose as in
laevifrons: inner lobe of maxillae c. as in
laevifrons and latipes, with hook from inner-
apical angle; palpi less thick than in pre-
ceding species. Prothorax: width length
1.33; base/apex 0.97; liase/head 0.68; mar-
gins narrow, much interrupted anteriorly;
disc weakly irregularly convex, irregularly
coarsely punctate. Elytra: width elytra/
prothorax 1.44; striae impressed, punctulate;
intervals convex, irregularly 2-seriately
punctate, 3rd with special dorsal punc-
tures not surely identifiable, 8th not much
wider than 7th, irregularly in part 2-seri-
ately sparsely punctate. Legs: tarsi mod-
erately wide and flattened, but less so than
in preceding species (latipes); 4th hind-
tarsal segments wide, very deeply emargi-
nate. Measurements: length 16.7; width
5.3 mm.
Type. Holotype $ (M.C.Z., Type No.
31,524), from Maba Vy., Menyama, Mo-
232
BuUetiu Museum of Comparative Zoology, Vol. 137, No. 1
robe Dist., N-E. N. G. (L. Hastings); 1 i
paratype (Bishop Mus.), Oriomo River, 6
m, Feb. 13, 1964 ("H.C."), light trap; 1
9 paratype (M.C.Z.), Maprik, N-E. N. (;..
Oct. 14, 1957 (Gressitt), hght trap.
Notes. This species is simihir to the pre-
ceding {hiti])cs) l)ut has more prominent
genae and narrower tarsi. The genae are
less prominent than in dinjsocomes but lead
toward that species, while the labrum sug-
gests a relationship \\ ith Jatipcs.
HELLUOPAPUA n. gen.
Dia^no.sis. Form of lleUuonidius; labrum
wide, multisetose; ligula wide, slightly
emarginate, setose as in llcUuonid\ns\ inner
lobe maxillae strongly hooked on inner side
before apex; Sth elytral intervals e. wide as
7th, 2-seriately punctate with few or no
scattered punctures; tarsi slender, 4th hind-
tarsal segments shallowly emarginate; front
femora obtusely pronu'nent below near base;
i front tarsi without squamae; i copula-
tory organs as in Figure LSI.
De.seription. See description of only
known species, below.
Ti/})e speeies: H. toxopei, below.
Gencrie distri])ution. Known from a
shigle locality in West N. (i.
Notes. This new genus differs from Ilel-
hionidiiis in labrum multisetose; hook of
inner lobe of the maxillae subapieal (not
apical), and tarsi much more slend(>r with
4th hind-tarsal segments shallowly (not
deeply) emarginate. It differs from Cwi,^/-
dema in labrum multisetose and (Sth el\ tial
intervals much narrow(M- and 2-seriatel\
(not densely) ptinctatc. It fits no other
genus in Sloane's (1914, pp. 571-572) key.
And it differs from all previously known
Ilelluonini of Sloane's ( 1914, p. 570) "Aus-
tralian (irouj) in lacking sexual s((namae
on i front tarsi.
Helluopapua toxopei n. sp.
Deseiij)li()iL With characters of genus;
form as in I'igure 146; slend(>r, subparallel.
depressed; antennae and hind tarsus, Figure
186; black, appendages dark brown, 2
minute red spots on head posteriorly; sur-
face sparsely pubescent, moderately shining,
microsculpture faint and irregular on head
and pronotum, more distinct and e. isodia-
metric on elytra, and surface punctate as
described below. Head 0.90 width pro-
thorax; genae rounded, moderately promi-
nent; clypeus slightly sinuate-truncate, with
several setae each side; labrum wider than
long, wide in front, broadly sinuate each
side in front with apex obtusely angulate,
with several principal setae each side but
n(jne near middle; front irregularly convex,
deeply impressed each side and transversely
impressed anteriorly (individual character?),
irregularly punctate at sides and posteriorly;
mentum with strong triangular tooth and
side lobes long and pointed; ligula as de-
scribed for genus; palpi rather slender.
ProtJiorax: width length 1.38; base apex
0.78; base head 0.69; margins narrow, not
interrupted; disc weakly irregularly convex,
surface irregularly pvmctate. Elytra: width
elytra prothorax 1.52; striae impressed, not
punctulate; intervals slightK' comex. rather
sparsely 2-seriately punctate. Measure-
ments: length 22.5; w idth 6.7 mm.
Type, llolotype S (sex determined by
dissection) (Leiden Mus.) from Rattan
Camp, West N. G., 1200 m, Feb.-Mar.
1939 (Toxopeus); the t\'pe is uniciue.
Notes. Although this ,j is from the same
localit)' as the 9 type of lleUuo)\idius
Idlipes. the two specimens differ in so man\'
ways that the\' cannot be one spt>cii\s but
ha\(' to be assigned to different genera:
the two specimens differ in lonn. in genae.
in sha])e and setae ol labrum, in position
of hook of inner lobe of maxillae, and in
form of tarsi, and this is just a beginning of
the list of differences. A r(>\ision ol all Ntn\
(iuinean and Australian members of the
tribe, with mucli additional material, will
l^robabK' be nec-essary to decide the real
relationships of this new genus as well as of
the new species ol I lelluonidiits described
above.
The Carabid Beetles of New Guinea • Darlington 233
Genus HELLUOSOMA Castelnau
Castelnau liSttT, Notes on Australian Coleop., p.
20.
Sloane 1914, Proc. Linnean Soc. New South Wales
39, pp. 571, 585.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1581 (see for synonymy and additional refer-
ences ) .
Diopiosis. See preceding Key to Genera.
Description. None required here.
Type species. H. atrum Castelnau (be-
low ) .
Generic distribution. Tropical Australia
and New Guinea.
Notes. Only one species of this genus is
adequately known, although a second
species may exist in Australia ( Sloane 1914,
p. 586).
Helluosoma afrum Castelnau
Castelnau 1867, Notes on Australian Coleop., p. 21.
Sloane 1914, Proe. Linnean Soc. New South Wales
39, p. 586.
Csiki 1932, Coleop. Cat., Caraliidae, Harpalinae 7,
p. 1581 (see for synonymy and additional refer-
ences ) .
Description. None required here; note
form c. as in Helluoniditis; color black or
dark brown; genae prominent; labrum
wider than long, obtusely angulate, 4-setose;
ligula narrower than usual in tribe, nar-
rowed anteriorly, narrowly rounded at apex;
length c. 12.5-15.0 mm.
Type. From Rockhampton, Queensland,
Australia; present location unknown (not
found at Melbourne in 1958).
Occurrence in New Guinea. Papua: 3,
Rouku, Morehead R., Apr. 1962 {\\\ W.
Brandt, C.S.I.R.O.); 7, Port Moresby and
vicinity, various dates and collectors ( Dept.
Agr. Port Moresby; A.M.N.H.); 1, Bisia-
numu, Sogeri Subdistrict, c. 1600 ft. (485
m). Mar. 1955 (J. J. H. Szent-Ivany and J.
McAdam, Dept. Agr. Port Moresby).
Notes. The relatively narrow ligula is
diagnostic of this species in this tribe in
New Guinea. I find no significant dif-
ferences between specimens from Australia
and from New Guinea, although individual
variation occurs in both places.
Genus HELLUODEMA Castelnau
Castelnau 1867, Notes on Australian Coleop., p. 19.
Sloane 1914, Proc. Linnean Soc. New South Wales
39, pp. .571, 586.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1581 (see for .synonymy and additional refer-
ences ) .
Diapiosis. See preceding Key to Genera.
Description. None required here.
Type species. HeUuomorpha hatesi Thom-
son {= tinicolor Hope), of Australia (see
below ) .
Generic distribution. Eastern and north-
ern Australia, New Guinea.
Notes. Two species of this genus occur
in Australia, one of them extending to New
Guinea.
Helluodema un/co/or (Hope)
Hope 1842, Proc. Ent. Soc. London for 1842, p. 47
( Aeni<itna).
Sloane 1914, Proc. Linnean Soc. New South Wales
39, p. 587.
Csiki 1932, Coleop. Cat., Caral)idae, Harpalinae 7,
p. 1581 (see for synonymy and additional refer-
ences ) .
Description. None required here; note
fonn (Fig. 141) slender; genae prominently
rounded; prothorax only moderately (not
strongly) constricted before base; labrum
wider than long, obtusely angulate, 4-setose;
ligula broadly rounded, obtusely emargi-
nate at apex; length c. 13-15 mm.
Type. From Australia; present location
unknown.
Occurrence in New Guinea. Papua: 3,
Rouku, Morehead R., Apr. 1962 {W. W.
Brandt, C.S.I.R.O.). West N. G.: 2,
Merauke, sea level. Mar. 24, 28, 1955 ( L. D.
Brongersma, Leiden Mus.), evidently taken
in light trap.
Notes. This species occurs in eastern
Australia at least from northern New South
Wales to Cooktown. The New Guinean
specimens agree well with Australian ones.
Genus GIGADEMA Thomson
Thomson 1859, Arcana Naturae, p. 93.
Sloane 1914, Proc. Linnean Soc. New South Wales
39, pp. 572, 593.
234 Bulletin Museum of Comparative Zoology. Vol. 137, No. 1
Csiki 1932, Coleop. Cat., Caiabidae, Harpalinae 7,
p. 1582 (see for synonymy and additional refer-
ences ) .
Diapwsis. See preceding Kcij to Genera.
Deseription. None required here.
Type .species. G. titanum Thomson ( =
nocte Newman), of Australia.
Generic distribution. Australia; south-
ern New Guinea.
Notes. This is the principal genus of the
tribe in Australia. One of the 12 or more
Australian species is now recorded from a
single locality in southern New Guinea,
almost opposite the tip of the Cape York
Peninsula.
Gigadema maxiilare Sloane
Sloane 1914, Proc. Linnean Soc. New South Wales
39, pp. 595, 599.
Description. None required here; note
large size; prothorax constricted at base;
color dark; surface short-pubescent; labrum
c. long as wide, rounded, 2-setose; length
of Australian specimens 27-35, of New
Guinean specimen 32 mm.
Types. From tropical Queensland, Aus-
tralia: Townsville, Kuranda, Cooktown,
Princess Charlotte Bay. I here designate
as lectotype a specimen labeled "Cktn., Q.,
Olive, i " and "Giiiadcmo nuixillare SI., Id.
by T. G. Sloane"; in Sloane Coll., C.S.I.R.O.,
Canberra ( seen ) .
Occurrence in New Guinea. Papua: 1
9 , Rouku, Morehead R., Apr. 1962 (W. W.
Brandt, C.S.I.R.O.).
Notes. The Papuan 9 agrees with Aus-
tralian examples of maxillare in nonsexual
characters, but a ^ is needed to confirm
the identification. Another specimen ap-
parently of nuixillare, but also a 9 , is
before me from Mona Is., Torres Straits
(J. VV. Schomberg, S. Australian Mus.).
Tribe BRACHININI
Brachijitiiii CIsiki 1932, (Coleop. Cat., (;aral)ida(\
Harpalinae 7, p. 1593 (sec for synonymy and
additional references ) .
Jedlicka 1963, Ent. .AbhanilliniHcn 28, p. 524.
Bnichinidac Jearniel 1942, Faime de France,
Coleop. Carabiques, I'art 2. p. 1102.
1949, Coleop. Carabiques de la Region
Malgache, Part 3, p. 1079.
BrachUdnac Basilewsky 1953, Exploration Pare
National I'Upemba, Fasc. 10, Carabidae, p. 235.
Habu 1967, Fauna Japonica, Carabidae, Trun-
catipennes Group, p. 280.
The beetles of this tribe are bombardiers
(but are not the only Carabidae that "shoot"
repellents) and are well known to most
entomologists in most parts of the world.
Their form is characteristic, and identifica-
tion is confirmed by presence of S visible
ventral abdominal segments.
Two genera of the tribe are very widely
distributed, and these are the only genera
that reach New Guinea. Fhcropsophus,
which occurs (discontinuously ) in all prin-
cipal tropical regions, has 6 New Guinean
species of which 5 are endemic and 1 shared
with Australia. Brachinus, which is almost
worldwide except that it does not reach
Australia, has 1 New Guinean species which
is endemic but which is the easternmost
member of an Oriental species group.
Key to Geneh.a of Br.\chixixi of New Guixe.\
1. Elytra with costae strong, distinct, and
separate to apex (p. 234) PJwropsopiuts
- Elytra with costae weaker and ])econnng
faint and in part vaguely connected before
ape-x (p. 239) Brarhiitus
Genus PHEROPSOPHUS Soiier
Solicr 1833, Ann. Soc. Ent. France 2, p. 161.
Csiki 19.32, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1595 (see for subgenera and ad(h'tional rcler-
ences ).
l\ira})hcropn(>]>h\is lliilxMitiuil 1911, Diiitsche \\\\[.
Zeitschritt for 1914. pp. 140. 442 (new s>-n()n-
yniy).
Csiki 1933, Coleop. Cat.. Carabidai'. Harpalinae 8,
p. 1604 (as subgenus of /'/icro/wo/^/n/.s).
Dia^^nosis. See preceding Key to Genera.
Description. None required here; see
Figures 147-151.
7'(//)<" species. Ol Pli(ropso))hu.s. Brachi-
nus senciialensis Dejean, oi Africa. Of
Paraplieropsoplius, P. intermedins Huben-
tlial {— verticalis Dejean) b\ present des-
ignation.
Generic distribution. All principal trop-
ical and s()m(- wann-teinperate regions of
The Carabid Beetles of New Guinea • Darlington 235
the world. In the Asia tic- Aiistrahan area
species are numerous in southeastern Asia
and the western Malay Archipelago, fewer
eastward, and only I variable species
reaches Australia.
Notes. The supposed "subgenus" Para-
pheropsophus was based on trivial char-
acters, primarily on the shape of the dark
mark between the eyes ( which is varialile )
supported by a supposedly characteristic
habitus which, according to Hubenthal, is
'ieichter zu erkennenden als zu beschrei-
benden." In my opinion all the 4 "species"
and 5 additional "varieties" listed in this
subgenus by Csiki (following Hubenthal)
are forms of a single species {verticals
Dejean) which is not worth subgeneric
separation from PJieropsophus.
The 6 species of Fheropsophus that occur
in New Guinea are ecologically differenti-
ated. P. verficalis is very common in a
variety of wet places and is winged. P.
amnicola has been found only on the banks
of large rivers (Markham and Sepik) and
is always winged. The other 4 species are
rare, local, flightless forms; 1 (cafiilus) is
known to occur in leaf-litter on the ground
in rain forest, and this is probably the habi-
tat of the others too.
Besides the 6 species of Phcropsophus
treated below, I have seen a single speci-
men of javanus Dejean (agnatus Chaudoir)
labeled as from New Guinea ("New Guinea,
Mimika R., A. F. R. Wollaston. 1911-229."
(British Mus.)). This conspicuous species
is common in the western Malay Archipel-
ago but has not been found in New Guinea
by recent collectors and has apparently not
been found in the Moluccas. I think the
specimen in question is probably wrongly
labeled. New Guinea should be deleted
from the range of the species as given by
Csiki (p. 1601) and others.
Key to Species of Fheropsophus of New Guinea
1. Elytra each with 8 costae c. equally promi-
nent 2
- Elytra with odd costae more prominent than
even ones at least at base ,5
2. Strictly bicolored: head and prothorax yellow
or reddish yellow without dark marks, elytra
dark without pale marks (p. 235) _ amnicola
- Not thus bicolored: all or part of pronotum
and part of head dark, elytra dark with or
without pale marks 3
3. Front yellow with isolated usually V-shaped
dark mark between eyes; inner wings large
and folded (p. 236) verticalis
- Posterior half or more of head dark; inner
wings vestigial 4
4. Head bicolored, yellow anteriorly, dark pos-
teriorly; prothorax wider than long at middle
(by measurement); femora not or only
minutely black-tipped; length 8.5-12.5 mm
(p. 237) aptim^iuorpJuis
- Head dark with small V-shaped reddish mark
on front; prothorax as long ( at middle ) as
wide and appearing longer; femora con-
spicuously black-tipped; length c. 18 mm (p.
237) pedes
5. Pronotum (sparsely) pimctate, much rough-
ened at base and apex; length c. 15-16 mm
(p. 238) catuht.s
- Pronotum \irtually impmictate, scarcely
roughened; length 20.5 mm (p. 238) __ cants
Fheropsophus amnicola n. sp.
Description. With characters of genus;
form (Fig. 147) c. as of verticalis; head
and pronotum yellow without dark marks,
elytra black without pale marks, append-
ages yellow. Head 0.92 and 0.93 width
prothorax. Prothorax subcordate; width/
length 1.14 and 1.13; base/apex 0.94 and
1.01; sides broadly arcuate anteriorly,
broadly sinuate before c. right posterior
angles; margins very narrow; disc convex,
irregularly subpunctate and punctulate.
Elytra moderately narrowed anteriorly;
width elytra/ prothorax 1.64 and 1.66; each
elytron with 8 well defined costae (in-
cluding raised suture), the costae equally
elevated and equally prominent at base;
surface of costae finely microreticulate, in-
tercostal intervals longitudinally roughened.
Inner icings full. Measurements: length c.
8.5-15.0; width 3.2-5.3 mm.
Types. Holotype c^ (M.C.Z., Type No.
31,525) and 34 paratvpes from vie. Nadzab,
N-E. N. G., July 1944 (Darhngton); and
1 paratype. Main R., Sepik, N-E. N. G.,
Feb. 1965 (R. Hornabrook).
Measured specimens. The S holotype and
1 9 paratype from Nadzab.
236 Bulletin Museum of Comparative Zoolop^y, Vol. 137, No. 1
Notes. The preceding description con-
tains all the characters that now seem worth
specifying for this new species, which is
distinguished from ceiiicaJis primarily by
color. (Jolor, pioperly understood and with
allowance for variation, is in fact specific
in this genus. The geographic and ecologic
restriction of amtiicola. as compared with
verticalis\ is another indication that the two
species are fully distinct. My specimens
were all taken on thi' banks of the Markham
R., and none was found in anv other situa-
tion.
Pheropsophus verticalis Dejean
Dfjfan 1825, Spitit-s (ieneral Coleop. 1, p. 302.
Csiki 19.33, Coleop. Cat., Caral)idae, Harpalinae
8, p. 1604 (see for Australian "varieties" and
additional references ) .
atistrali.s Castelnau 1867, Notes on Australian
Coleop., p. 23 (new synonymy).
papucmis Maeleay 1876, Proc. Linnean Soc. New
South Wales 1, p. 166 (new synonymy).
Heller 1910, AbhandhuiKen und Berichte Zool.
VIus. Dresden 13, No. 3, p. 7.
luaclcayi Sloane 1894, Proe. Linnean Soe. New
Soutli Wales (2) 9, p. 4.53 (new synonymy).
haliothonix Heller 1910, Abhandlunuen und
Berichte Zool. Mus. Dresden 13, No. 3, p. 6
( new synonymy ) .
intermt'dius Iluhentiial 1914, Deutsclit' Ent.
Zeitschrift for 1914, p. 440 (new synonymy).
Deseription. None required here. This is
the only New Ciuinean Plwropsophiis with
an isolated black (or brown) frontal spot.
See also under genus (above) and Notes
(below); length 8.5-16.5 mm.
Types. Of verticalis Dejean, from
"Nouvelle Tlollande" (= Australia ); in
Oberthiir Coll., Paris Mus. Of (tiislralis
Castelnau, from Hockhampton, Queensland,
Australia; present location unknown. Of
ixipnemis Maeleay, from Katow , Papua;
presumably in Maeleay Mus., Sydney. Of
nuirlediji Sloane, from King's Sound, NW.
Australia; lectotype not designated. Of
haliolliorax Heller, from l''iiischhafen, N-K.
N. (i.; in Dresden Mus. Of intermedins
Ilubenthal, from Neu Itrilain; in Hcrlin
Zool. Mus. (Of all these types, I have seen
only some cotxpes of niaeleai/i. )
Occurrence in New Cui)\ea. Coinmou
throughout New Guinea: 215 specimens;
most from low altitudes, only 2 individuals
from above 1000 m; common at Dobodura.
Notes. The synonymy proposed abo\'e is
based not on comparison of types but on
examination of much material from many
localities in Australia as well as New
Ckiinea, New Britain, and some other islands.
As a result of it I have concluded that, in
the area in question, all the Fhcropsophus
with an isolated dark frontal spot belong
to one variable species, verticalis Dejean,
which ranges over the whole of Australia
and New Guinea and extends to New
Britain, New Ireland, the Solomons, and
perhaps other islands.
In reaching this conclusion, I first con-
sidered the supposed separate northern
Australian species, maclcayi Sloane. Of it,
Sloane had only 3 specimens, which hap-
pened to be rather small ( 11.5-13 mm) and
to share some minor peculiarities of form
and markings including presence of yellow
shoulder spots and elytral fasciae of con-
stant shape. My material from subtropical
and tropical northern Australia shows that
these characters are in fact indixidual rather
than si)ecific. For example, 6 specimens
from the Blackall Range, in subtropical
South Queensland, \ary in size from 11.5
to 14.5 mm ( to apex of {>K'tra ) and vary
also considerabl) in lorm and somewhat in
markings, although none has shoulder
spots; and 4 specimens from mid-j:)eninsular
C>ape York (Co(mi and Silver Plains) are
large (e. 14.5-16.5 mm) and \ary in exact
form of prothorax, in prominence of humeri,
and in markings: e.g., 2 have and 2 have
not Ncllow shoulder spots.
The New Guinean individuals that I in-
clude in verticalis vary in details of form,
especially in prominiMu-e of humeri. 4'he\-
\ary in si/.r Irom e. S.5 to 16.5 mm. And
tlu'v N'ary in markings: the trans\(M-se usu-
all\ \'-shaiied dark mark between the eyes
is relatixcK constant but is soiiictimes
slighth extentled posteriori) ; the pronotum
\aries Irom wholly dark to broadl\- reddish
Ncllow with ()nl\- the margins dark (inter-
The Carabid Beetles of New Guinea • Darlington 237
mediates are common ) ; and the elytra vary
from \\'holly dark to conspicuously marked,
with median fascia often present but vari-
able (but rarely large), shoulder spots
sometimes present (distinct in only 2 in-
dividuals, from S. Highlands and Popon-
detta, and vestigial in a few other individ-
uals), and apices sometimes with (variable)
yellow margins. Although much of this
variation is surely individual, some of it is
or may be geographic. For example. New
Guinean specimens usually have the elytra
less heavily spotted than Australian speci-
mens, although extremes overlap. But I
think nothing is to be gained by recognizing
subspecies now. The variations of this
species should first be analyzed statistically,
in detail, using series of specimens from
exact localities, not just the New Guinean
against Australian specimens. This will be
third stage taxonomy (see Part I of my
■'Garabid Beetles of New Guinea," p. 329),
far beyond what I can attempt now.
The wings of vcrticalis are fully de-
veloped, or at least large enough to be
strongly folded at apex, in all my Australian
specimens and all New Guinean specimens
that I now assign to this species. However,
occasional short-winged individuals occur
that may prove to be mutants of veiticalis
although I am tentatively treating them as
a separate species, ciptinomorphtis Heller
( below ) .
P. vcrticalis is common in a variety of
wet places. Although all individuals are
winged, they may not often fly and are
not often taken in light traps. Observations
on their flight would be interesting.
The great variation in size of adults sug-
gests that the larvae may be parasitoid, per-
haps feeding on pupae of other beetles, as
some other members of the tribe are known
to do.
Pheropsophus apfinomorphus Heller
baliothorax var. apfiiH>iuori)}ui.s Heller 1910,
Abhandlun^en unci Berichte Zool. Mus. Dresden
13, No. 3, p. 7.
Description. Form as in Figure 148;
similar to vcrticalis (above) but elytra
more narrowed to base; head bicolored,
yellow anteriorly, dark posteriorly; elytra
not marked (in the few specimens seen);
inner wings vestigial, reduced to thin strips
less than Vj long as elytra; length (to apex
of elytra) c. 8.5-12.5 mm.
Type. From New Guinea, exact locality
not given, but altitude stated as 120 m; in
Dresden Mus. (not seen).
Occurrence in Netc Gtiinca. N-E. N. G.:
1, Aitape, Aug. 1944 (Darlington). West
N. G.: 1, Waris, S. of Hollandia, 450-500
m, Aug. 16-23, 1959 (T. G. Maa, Bishop
Mus.); 1, Maffin Bay, Aug. 1944 (Darling-
ton); 1, same locality, June 15, 1944 (E. S.
Ross, Galifornia Acad. ) .
Notes. Heller does not say whether the
type is winged, but the 4 specimens listed
above answer his description in color of
head (which is diagnostic), and their form,
with narrowed humeri, does recall A))tinus.
If it were not for the different head
marking, I would consider my specimens of
aptinomorphus to be short- winged mutants
of vcrticalis. The distribution of the short-
winged indi\iduals, widely scattered and
occurring with vcrticalis (which I have
from Waris and Maffin Bay and which
probably occurs at Aitape too), would be
consistent with their being mutants. But
if they are mutants, then the mutant gene
apparently must modify color of head as
well as length of wings and form of elytra,
and I do not dare assume that this is the
case without further evidence.
Pheropsophus pedes n. sp.
Description. With characters of genus;
form as in Figure 149, with relatively long
narrow prothorax and elytra ample but
strongly narrowed to base; black, head with
irregular red marks including U-shaped
mark between eyes, appendages reddish,
femora broadly tipped with black. Head
0.96 width prothorax; eyes normal; 1st
antennal segments swollen; labrum semi-
circular; front 2-impressed anteriorly, with
surface finelv c. isodiametricallv reticulate.
238
BuUciin Museum of Comparotivc Zoolomj, Vol. 137, No. 1
Profhomx long; width/length 1.01; base '
apex 1.06; margins narrow, each with 1
principal seta behind middle of length; disc
moderately conxex, ^^'ith scattered punc-
tures (the punctures with setae as usual in
genus), strongly roughened at base, less so
apically, and with disc finely irregularly
microreticulate. Elytra: width elytra/pro-
thorax 1.95; each elytron with 8 costae (in-
cluding raised suture) c. equalh' dexeloped;
siuface of costae finely irregularh' micro-
reticulate, intercostal intervals longitu-
dinally roughened. Inner aings evidently
atrophied. Measurements: length LS; width
6.5 mm.
Ti/pe. Ilolotype 9 (Bishop Mus.) from
Bomberi, Vogelkop, West N. G., 700-900
m, June 7, 1959 (T. C. Maa); the type is
imique.
Notes. I do not know the relationships
of this obviously distinct species. Char-
acters distinguishing it from other New
Ouinean Plier(>})so})1ius are given in th(> pre-
ceding Key tu Speeies.
Pheropsophus cafulus n. sp.
Deseription. With characters of genus;
form as in Figure 150, with elytra strongly
narrowed to base; brownish black, head
variably red-marked anteriorly, appendages
reddish'. Head 0.91 and 0.90 width pro-
thorax; eyes moderate; 1st antennal seg-
ments slightly swollen; labrmn transverse,
slightly prominent at middle; front 2-im-
pressed anteriorly, with surface finely ir-
regularly microreticulate, roughened jios-
teriorh. Prothorax (juadrate-subcordate;
width length l.OI and 1.07; ba.se/apex 1.11
and 1.13; sides weakly arcuate anteriorly,
broadly sinuate before basal angles; latter
acute except bhmted, slightK produced
posteriorly; margins moderate, each with
one principal seta behind middle of k'ugth;
disc irregularly convex, irregularh rough-
ened especially posteriorly and anteriorly,
and surface also with scatteri'd punctures
(with hairs as usual) and irregularly faintly
microreticulate. Elytra: width elytra pro-
thorax 1.72 and 1.74; each cKtron with S
costae (including raised suture), even
costae stronger than odd ones and reaching
base; surface of costae faintly finely micro-
reticulate, intercostal intervals finely longi-
tudinally roughened, and alternate intervals
each with row of widely spaced setae.
Inner wings reduced to vestiges that hardly
extend beyond edge of metathorax. Mea-
surements: length e. 15-16; width 5.0-5.6
mm.
Types. Holotype i (M.C.Z., Type No.
31,526) and 2 9 9 paratypes all from
Dobodura, Papua, Mar.-July 1944 (Dar-
lington ) .
Measured specimens. The i holotype and
1 9 paratype.
Notes. I do not know the relationships of
this very distinct species, which is quite
different from any of the preceding ones
although closely related to the following
(canis). The beetles were taken among
dead leaves on the ground in rain forest.
Pheropsophus canis n. sp.
Description. With characters of genus;
form as in Figure 151, with elytra strongly
narrowed to base; brownish black, head red
anteriorly dark posteriori)-, pronotum with
faint reddish marks, legs reddish yellow,
antennae brown. Head 0.S7 width pro-
thorax; eyes moderate; 1st antennal seg-
ments slightly swollen; labrum transverse,
with margin broadh' rounded; front weakh'
2-impressed, closely irregularly microreticu-
late, roughened posteriorly. Prothorax nar-
rowK' subcordale; width length 1.07; base '
apex 1.06; sides broadU' arcuate anteriorh,
broadly sinuate before slightly obtuse,
slightly blunted, posterior angles; latter not
produced posteriorly; margins moderate,
each with 1 principal s(>ta behind middle;
disc moderately conxcx. weakh roughened
posteriorly but otherwise uearK' smooth,
surlace w ith Wwc irregular reticulate" micro-
sculpture. Elytra: width eKtra i^rothorax
l.(S(); each eKtron with 5 conspicuous costae
(nos. 1, 3, 5, 7, (S), the intermediate costae
(nos. 2, 4, 6) weak or obsolete; surface^ of
costae lineK' irreirularK microreticulate. iu-
The Carabid Beetles of New Guinea • Darlington 239
tercostal intervals finely roughened; even
intervals probably with widely spaced setae,
but latter in part broken oft or missing.
Inner wings evidently vestigial. Measure-
ments: length c. 20.5; width 7.6 mm.
Type. Holotype 9 (Hawaiian Sugar
Planters Association) from Koitaki, Papua,
1500 ft. (c. 460 m), Nov.-Dec. 192(S ( Pem-
berton Coll.); the type is unique.
Notes. This flightless species is probably
a geographic representative of the pre-
ceding (eatuhis) but is larger, slightly dif-
ferent in proportions, with antennae darker,
pronotum much less roughened, and even
intervals of elytra much more reduced.
Other, related forms of this flightless group
are to be expected elsewhere in New
Guinea.
Genus BRACHINUS Weber
Weljer 1801, Observationes Entomologicae, p. 22.
Bidclujnus Auct. including Csiki 1933, Coleop.
Cat., Carabidae, Haipaliiuu' (S, p. 1606 (see for
additional references ) .
Diagnosis. See preceding Key to Genera.
Description. None required here.
Tyi)e species. Carahus crepitans Lin-
naeus, of Europe.
Generic distribution. Most of the world,
except Australia. In the Indo-Australian
area, many diverse species occur in India,
etc.; fewer, in the western part of the Malay
Archipelago; and a single species occurs
in New Guinea.
Brachinus papua n. sp.
Description. Form as in Figure 152; dark
brown or brownish black, head red anteri-
orly or with 2 red spots between eyes,
pronotum sometimes vaguely reddish, ap-
pendages reddish with tibiae, tarsi, and
apices of femora darker brown; dull, entire
upper surface with fine, c. isodiametric but
irregular reticulate microsculpture, and
much of upper surface with inconspicuous
fine pubescence, often in part rubbed away,
and perhaps missing on front, part of pro-
thoracic disc, and disc of elytra. Head 1.01
and 0.99 width prothorax; eyes moderate,
genae oblique, setose; front longitudinally
impressed each side, nearly smooth or
slightly roughened and punctulate ( vari-
able), with head more roughened posteri-
orly. Prothorax subcordate (exact form
variable); width/length 1.07 and 1.07; base/
apex 1.03 and 1.01; margins moderate, each
with apparently 1 principal seta near middle
of length; disc weakly convex, slightly finely
transversely wrinkled, irregularly punctu-
late, slightly longitudinally roughened at
base and apex. Elytra ample; width elytra/
prothorax 2.34 and 2.33; intervals slightly
raised but not costate. Inner icings full.
Measurements: length 17.5-19.0; width
7.3-8.0 mm.
Types. Holotype c5 (M.C.Z., Type No.
31,527) from Hollandia, West N. G., 250
ft.. May 1945 (H. Hoogstraal); 1 paratype,
same locality, Apr. 1945 (B. Malkin,
U.S.N.M.); 2 paratypes, Tanahmerah, Res.
Boven Digoel, West N. G., Feb. 1958
( R. T. Simon Thomas ) ; and 1 paratype,
Fenichel (Hungarian National Mus.).
Measured specimens. The S holotype and
1 9 paratype from Tanahmerah.
Notes. This, the first known New
Guinean Brachinus, may represent higut-
ticeps Chaudoir of Java, etc., but papua is
larger than my specimens of bigutticeps,
with less sharply bicolored legs and less
roughened pronotum. Further study is
needed to clarify the relationships of these
and related species in the Malay Archipel-
ago.
Tribe PSEUDOMORPHINI
Csiki 1933, Coleop. Cat., Carabidae, Harpalinae 8,
p. 1634 (see for synonymy and additional refer-
ences ) .
Pseiidomorphidae Auct. including Notman 1925,
Proc. United States National Mus. 67, Art. 14,
p. 1.
Pseudomorphini (Figs. 153-159) do not
look like Carabidae but superficially re-
semble dytiscids or scolytids or Crypto-
cep]}alus-\ike chrysomelids. They are nu-
merous onlv in Australia; a few small
species occur in New Guinea; a species of
240 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
the Australian and New Guinean genus
Adelotopus has lieen found m Java; one
genus, Cnjptocephalomorpha, ranges from
New Guinea aeross the Malay Archipelago
to the SE. corner of Asia; and a supposed
endemic genus is localized on New Cale-
donia. Outside this area the tribe contains
only a single genus, Fseudomorpha, con-
fined to the Americas and ranging from
-southern United States to Brazil and
Argentina/
Although I have only 15 specimens of
this tribe from New Guinea, they include 3
genera and 7 species, and all the species are
different from the single pseudomorphine
{Adelotopus papuanus) previously known
from the island. ( An ^^ Adelotopus sp.^' listed
from New (Guinea by Heller, in Abhand-
lungen und Berichte Zool. Mus. Dresden
13, 1910, No. 3, p. 4, has not been identi-
fied.) Because my material is very limited
and because the species are well defined
by easily seen characters, I shall treat the
members of this tribe rather superficially,
leaving dissection of the mouthparts, etc.,
to the next reviser of the tribe as a whole.
Notman's ( 1925 ) keys to the Australian
species of this tribe are very useful but,
because they are based largely on old de-
scrij^jtions rather than on specimens, the\'
should be used with caution.
Most Australian Pseudomorphini live on
the tnuiks of trees, especially on the shaggy
trunks of Euealyplus. They are v(My active,
winged beetles. I do not know tiie habits
ol the New Guinean h)rms.
' The lollowiiiji; notes arc necessary to justify
my siininiar\ of [\\c distrihiition ot this zoot^t'o-
.Uraphically iiilcrcstinu tribe. The African Ihj-
ilroporoDiorplui has only 1 entire \entral abdominal
sesnients; it is not a pscucloniorphine but probabK
a harpaline. Paus.sotwpus is Australian and jirob-
ably does not occur on Batchian Is. ( \otnian
1925, p. 5, footnote; conlirnied 1)\ nie at the
British Mus. in 1948). And SilplioiiioiplKi
cntiaroidcs (Newman), listed from Oceania 1)\
Csiki (1933, p. 1639), is probably really Aus-
tralian; Newman kIvcs no localit\' except "Its
habitat is 3753," but lie refers to the inseit as
''this jircttx' antipodean.
Key to Gener.\ of Pseudomorphini of
New Guinea
1. Eyes superior in position, not interrupting
lateral margins of heatl (p. 240) _ _ Adelotopus
- Eyes lateral in position, broadly interrupting
lateral margins of head __ 2
2. Form wider, more depressed; head not
strongK deflexed, labrum and mandibles
\isible from in front (p. 242) __ SpliaJIoiuorpha
- Form subc\lindric; head sti"ongl\ deflexed,
labrum and mantlibles ( except sometimes
their tips ) not \ isible from in front ( p.
242 ) Cnjptocephdiontorplia
Genus ADELOTOPUS Hope
Hope 1834, Trans. Ent. Soc. London 1, p. 11.
Notman 1925, Proc. United States National Mus.
67, Art. 14, pp. 5, 6.
Csiki 1933, Coleop. Cat., Carabidae, Harpalinae 8,
p. 1634 (see for additional references).
Diciii^nosis. See preceding Key to Genera.
Description. None required here.
Type species. A. '^yrinoidcs Hope, of
Australia.
Generic distribution. Australia including
Tasniania (many species). New Guinea
( 4 species ) , and Java ( 1 species ) .
Notes. A. jaeohsoni Ritsema ( 1909, Notes
Leiden Mus. 31, p. 255) of Java evidently
realK' is an Adelotopus, not a Cryptoeephalo-
morpha, for Ritsema knew both genera.
Key to SrEf:]Ks of Adei.otoi'I s of New GnNF..\
!. Piceous, with narrow reddish tianslucent
margins i)ut otherwise unmarki'd; l